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Britisb museum (Ratiiral l)lstorp). 

This IS No. 25. of 25 copies of 

the " Catalogue of Chiroptera " {Second 
Edition), Vol. 1, printed on special 



C H I R P T E R A 








Volume I : MEGACHIROPTER:^^'^ '^'^^'^^^ 

■ r 





DVLAV & Co.. Ltd., 37 SOHO 8Q0ARE, W. ; 




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The Cntalogjiie ■oi Chiroptem in the British Museum by Dr. G. 
E. Dobson, published b}- the Trustees in 1878, proved to be, as 
was expected, a work of great utility to zoologists, the systematic 
arrangemeut of these animals having bceu previously in a state 
of the utmost confusion. The Catalogue was therefore generally 
welcomed, and as a consequence soon ran out of print and became 
almost unobtainaMe. Iforeover, the stimulus it gave to the study 
of Bats rapidly tended to render it obsolete, and a new edition has 
therefore long been a desideratum. 

But owing to the difficulties of the subject and the necessity that 
the writer should be able to give his whole time to the work, 
undistracted by official duties, it has not previously been possible 
to arrange for tlie publication of a second edition. 

Two or three years ago, however, the Trustees were fortunately 
able to secure the services of Dr. Ivnud Andersen, who had already 
made a special study of Bats, and who has been able to devote 
himself uninterruptedly to the preparation of the present woik. 

This Kdition is in reality a completely new and (irigiiial Mono- 
grajdi of the Order Cbirnptera, ior materiids. metliods of work, and 
ideas on si>ecies have all so radically ch:ingcd since 1878, that 
notliing remains of the first edition — good as that was for its 
date— but the title. 

So great is the increase in the general knowledge of the subject 
and also in the material examined and described, that whereas in 


1878 one volume was fouud sufficient for the whole of the Order, 
rather less than one fifth of it being devoted to the Mega- 
chiroptera, a volume of somewhat greater size is now needed for 
the Megachiroptera only, while a corresponding enlargement will 
probably take place in the part dealing with the other groups of 
the Order. 

The Author indicates in his Preface that the number of specimens 
belonging to the Museum has greatly increased since 1878, while 
in addition he has been able to examine all the chief collections of 
Chiroptera in Europe, notably those of Paris, Berlin, and Leyden, 
and to borrow a large part of that of the U.S. National Museum at 
Washington. The thanks of the Trustees are due to the Authorities 
of the various Museums who have helped him in this respect. The 
■volume also' owes much to Mr. Oldfield Thomas, F.R.S., who 
has interested himself in its progress since its commencement, 
and has assisted the Author by discussing difficulties from time 
to time as they arose, and in other wars. 

British Museum (Natural History). 
February 22nd, 191--'. 


Keeper of Zoology. 

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SpECiMhw.s EXAMiXED. — III 1843, wheii J. E. Gray published his 
' List of the Specimens of Mammalia in the British Museum,' the 
Megachiroptera were represented in the National Collection by 
69 specimens. Thirty-five years later (Dobson's ' Catalogue of 
Chiroptera,' 1878) the number had increased to 425. After 
another period of thirty-four years (this Catalogue) the total 
reaches 1470. The skins enumerated in this volume amount to 
956, the specimens preserved in alcohol to 444, and the skulls to 
1228. Exactly half a century ago (Gerrard's 'Catalogue of the 
Bones of Mammalia in the British Museum,' 1862) the number of 
skulls was only 39 ; in 1878 it was 82. That the number of 
specimens is now three and a half times, but that of skulls fifteen 
times, greater than in 1878 is due to the fact that during the 
preparation of this Catalogue the skulls have been extracted from 
nearly all the old skins in which they had hitherto been left as 
well as from about 60 per cent, of the alcoholic specimens. In 
Dobson's time there were separate skulls of only half the number 
of species of Megachiroptera then in the Collection, whereas now 
every species and subspecies in the Museum, with one exception, 
{Pteropus arncnsis), is represented by at least one and often by a 
series of skulls. 

Besides the specimens preserved in the British Museum I have 
had for inspection a large number from other Collections, and 
during two visits (in 1907 and 1909) to the Museums of Leyden, 
Berlin, and Paris I had the privilege of going through nearly 
the whole of the series of Megachirojjtera in those Collections, 
so that the total number of specimens examined for the purpose of 
this Catalogue amounts to about 2400. 


Number of genera, species, ami subspecies. — The growth of our 
knowledge of the existing forms of Megachiroptera, since Linnean 
times, is shown in the subjoined table : — 


1758. Linno t . 

1810. E.Geoffroyt 2 . 

1825. Temiuinck{ 2 . 

1837. TemminckJ 5 . 

1867. Peters 1 9 . 

1870. Gray* 14 . 

1878, Dobson 10 . 

1899. Matsohie + 20 

1907. Millerl 30 

1912. This Catalogue 35 . 

Genera and 



Species. Forms. 


, 20 


49 . 

57 . 
. 70 
. 120 





Of the 228 forms described in this Catalogue 207 (91 per cent.) 
tire represented in the Collection, of the 35 genera 32 (91 per 
cent.). The following twenty-one forms are at this moment 
desiderata (the throe missing genera, all mouotypic, are marked 
with an asterisk) : — 

rteropus /ii/j)omelanus geminontm, 
p. 1C6. 

„ pallidus, p. 136. 

„ pwmilus, p. 757. 

,, ualanus, p. 177. 

„ mariantius, p. 178. 

„ va/iil-ore/isi^, p. 184. 
"" „ aldahmisis, p. 213. 
■i; '„ tubcrculatus, p. 309. 
_ J ,, vampj/rua plidon, p. 353. 

,, macrotU, p. 396. 
Aexrodon lucifer, p. 432. 

* Plcroies andiiota, p. 486. 
Epomoplorus pousarguesi, p. 543. 
Myoiigoteris {I'hygetis) brachyce- 

■ph(da, p. 582. 
Cynoplerus bracliyotis miuutus, 
p. 625. 
,, vuij&r, p. 029. 

,, lyrinceps, p. (i33. 

* Pieuochincsjagori, p. 645. 

* Chironax melanocephalus, p. 660. 
Syconycteris anstralis, p. 781. 
Noiopieris neocaledonioa, p. 799. 

All forms, including those not represented iu the Collection, have 
been examined by me. 

t All Chiroptera were iu the tenth edition of the ' Systeiua Natura; ' referred 
to one genus, Vefperfilw. 

\ E. Geofiroy, ' Description des Roussettes et des Cephalotes, deux nouveaux 
genres de la faiuille des Chauve-souris ' ; Ann. Mus. d'Hist. Nat. Paris, xv. 
pp. 86-108 (1810). 

Teraniinck, ' Monographies de Mainmalogie,' i. pp. 166-204 (1825). 

Temminck, vp. cit. ii. pp. 49-112 (1837). 

Peters, " Ueber die Flederlmnde, Ptcrop't, und insbesondere iiber die Arten 
der Gattung Pterrpus s. s." ; MB. Ak. Bedin, 18G7, pp. 319-333, 86.>-872. 

Gray, 'Catalogue of the Monkeys, Lemurs, and Fruit-eating Bats in the 
Collection of the British Museum' (1870). 

Matschie, ' Die Megachiroptera des Berliner Museums fiir Naturkunde ' 

Miller, " The Families and Genera of Bats " ; U.S. National Museum, 
Bulletin 57 (1907). 

authdks teeface. 

Types of valid names (liuti/pes).— The types of the 228 species 
and subspecies of Megachiroptera are distributed as follows: — 

Kidolon (3 forms) 1 

H(),I^<;ttus (U) I 9 

lioiK'ia (\) ' 

Pteiopus (103) I 49 

Acerodoii (9) I 5 

Pteralopex (2) j 2 

Styloctcnium (1) 1 

Dobsonia (18) j 8 

Harpi-ionycteris (1) ... 1 

Plerotes (1) I 

Epoiuops (4) 1 

Hvpsigiiatlms (1) I 

Epoinopliorus (9) | 3 

Mici'opteropus (1) 

Nanonyctoris (1) 

Scotouycteris (1) 

C'jwiuycteris (1) .. 

Myouycteris (4) ... 

Cyhopterus (16) ... 

Ptciiochinis (1) ... 

Me;ja;rops (1) 

I Dyacopterus (!)■ ... 
I Halionycteris (1) ... 

! Chironax (1) 

j Tliooptevus (1) 

I I'oiithetor (1) 

j Splia;rias (1) 

I Xyctimeiic (13) ... 

j Koiiycteris (3) 

} Mcgaloplossus (1) 

Macioglossus (6)... 
' Syconycteris (7) ... 

Mi'loiiycteiis (1) ... 
] Nesonycteris (1) ... 

Notopteris (2) 

:= t3 

^ 13 



Not traced, 1. 
Calcutta, 2 ; Li,-,boii, 1. 

Caloitta, 1; Cambridge (Mass.), 1; 
Copenhagen, 1 ; Genoa, 3 ; Sydney, 
1 ; ? Zi-ka-wei, 1 ; private posses- 
sion, 1 ; not traced, 5. 

Chicago, 1 ; Dorpat, 1. 

Brisbane, 1 ; ? Zi-ka-wei, 1. 

liisbon, 1. 

Stuttgart, 1. 
Not traced. 

1 ; Stockhohti, 1 

Lisbon, 1. 

Calcutta, 1 ; Vienna, 1; not traced, 1. 


Not traced, 1. 
Calcutta, 1. 
Not traced, 1. 

yummed up, this gives the following totals: — British MuseuiQt 
118, or more than half the number of all cutypcs; Berlin Mu- 
seum, 25 ; Leyden, 22; Paris, 17; U.S. Natioual, 15 ; Calcutta, 5; 
Lisbou, 4; Genoa, 3; two each in the Philadelphia and Zi-ka-wei (?) 
Museums; one each in the Brisbane, Cambridge (^Massachusetts), 
Chicago, Copenhagen, Dorpat, Hamburg, Stockholm, Stuttgart, 
Sydney, and Vienna Museums ; one {Ptcropas lijtleri) is said to be. 
in private possession ; while ten have not been traced (moat of 


these, if not all, are almost certainly not in existence), viz. Eiclohn 
heJvxim (once in the Mnseum Leverianum), Pteropus hypomelanus 
tomesi (once in the possession of Mr. L. L. Dillwyn), Pteroims 
suhniger (Cabinet lleaumur), Pteropvs rufus (Paris Museum), 
Pteropus nir/er (ancient Royal Cabinet, Paris), Pteropus polio- 
ceplialus (based on specimens in Leyden, Paris, and London, none 
of which can now be traced with certainty), Micropiteropus p>usillus 
(cotypes once in the Tomes Collection and Paris Museum), Cijno- 
pterus sphinx (ancient " Naturhistorie-Selskab," Copenhagen), 
Nyciimene cephalot€s (described by Pallas), and Macror/lossus 
minimus (Paris Museum). — That the sum total (234) is a little 
in excess of the number of recognized forms (228) is due to the 
fact that in a few cases the cotypes of one form are preserved in 
different collections. 

The 228 forms have been described by the following fifty-seven 
writers (those, seventeen in number, who have described eutypes 
preserved in the I3ritish Museum are marked with an asterisk) : — 

Allen (G. M.), 1. 
Allen (H.), 1. 

* Andersen, 57. 

* Anderson & de Win- 

ton, 1. 
' Blyth, 1. 

* Bocage, 'i. 
Briinnich, 1. 
Desniarest, 2. 
De Vis, 1. 

* Dobson, 13. 
Elliot, 1. 
Eschscboltz, 1. 
Geoffroy (E.), 7. 

* Gould, 1. 

* Gray, 16. 
Hallowell, 1. 
Heude, 2. 
Heuglin, 1. 

* Hodgson, 1. 
Hombron & Jacquijiot, 

Jentink, 3. 
Kelaart, 1. 
Kerr, 3. 

* Lay, 1. 

* Linne, 1. 

* MacGillivray, 1. 
Mason, 1. 
Matscbie, 8. 
Mearns, 2. 
Miller, 10. 
Milne-Edwards, 1. 
Miiller (S.), 1. 

* Nicoll, 1. 

* Ogilby, 1. 
Pageustecher, 1. 
Pallas, 1. 

Peale, 1. 
Pelers, 20. 
Peters & Doria, 1. 
Quoy & Gaimard, 3. 
Ramsay, 1. 
Scblegel, I. 

* Sclater (P. L.), 1. 
Seabra, 1. 

* Smith (A.), 1. 
Sundevall, 1. 
Teinminck, 15. 

* Thomas, 24. 

* Thomas & Wroughton, 


Tomes, 1. 
Tronessart, 2. 
True, 1. 
Vabl, 1. 
Zelebor, 1. 

The types of 185 forms have been examined by me, and of ten 
other forms I have had paratypes for inspection. With regard to 
the remaining thirty-three forms, I have seen topotypes of nineteen, 
of two types I have had photographs of the skull and dentition, 
and two have been examined for me by others. This leaves only 
ten forms, but of six of these the types are, either certainly or 
probably, not in existence, leaving finally the following four, 
viz. Pteropus papuanus, Dobsonia pannietensis, Epomops dobsoni, 
and Megalo(jlossus xvoermanni, but in none of these four cases is 
there any doubt as to the identification of the species. 


Types of synonyms {Hypotypes). — 127 names (variants not 
counted) occur iu the synonymy of the recognized forms of 
Megachiroptera. Eight are noraina nuda, and the types of 
thirty-nine names have not been traced. The remaining eighty 
hypotypes are distributed as follows : — 

British Museum, 38; Berlin, 10; Leyden, 7; U.S. National 
Museum, 6 ; Paris, 5 ; Zi-ka-wei (?), 3 ; two each in the Museums 
of Frankfort-on-the-Main, Lisbon, Philadelphia, and Sydney ; and 
one each in Calcutta, Dresden, Stockholm, and Vienna ; while one 
is probablj* in private possession. 

63 of these SO types have been examined by me, while of two 
other forms I have seen paratypes. In thirteen other cases I have 
either had photographs of the skull and dentition of the types, or 
the t3-pe8 have been examined for me by local zoologists, or I have 
seen topotypes. The remaining two names are Pteropus masMrinus 
(= Ft. rodricensis) and Odontonycteris meyeri (= Macroglossus 

Tlie Catalogue. — Nominally this is a second edition of Dobson's 
' Catalogue of the Chiroptera in the British Museum ' (1878^ ; in 
reality it is an independent work, except of course in so far as 
every work of this kind must be based on its predecessors. The 
descriptions are new, the technical names are fixed in strict 
accordance with the principle of priority, the synonymy is in most 
cases ■worked out on the basis of an examination of the types and 
paratypes, and all references to literature have been compiled by 
myself. It has been my object on the one hand to make the 
descriptions of the genera, species, and subspecies reasonably com- 
plete, on the other hand to avoid repetitions as far as possible. For 
the latter reason I have often preferred to give a differential rather 
than a full description, that is, I have confined the description of a 
given form chiefly to those characters by which it differs from its 
nearest fully described relatives. To render the Catalogue easier 
for reference the descriptions are, whenever required, subdivided 
into specially headed paragraphs (Diagnosis ; Skull ; Dentition • 
Palate-ridges ; External characters ; Sexual differentiation ; Speci-' 
mens examined ; liange ; Habits ; Affinities ; History in literature ; 
Type, Nomenclature, and Synonymy, »tc.). Detailed measurements 
are given of the skull aod external dimensions, and as a rule also 
of the premolars and molajs, of every form described. 


Ill the Introduction to this volume (pp. xvii-ci) I have given a 
summary of the general characters of the Megachiroptera, of the 
jirincipal variations within the Suborder of the cranial, dental, and 
external characters, of the mutual affinities of the genera, of the 
geographical distribution of the genera, species, and subspecies, 
and finally, a sj-nopsis of the more important differential characters 
of the subfamilies and geuera, and an artificial " key " to the 

The illustrations (80 in number ; see list p. 835) are original, 
with two exceptions (fig. 32, p. 4!)0, and fig. 51, p. 643), and 
figures are given of the skull and dentition of all genera and 
subgenera, except one (Chironax, p. 658). 

As this is the first Vertebrate Catalogue in which the register 
numbers of the specimens have been published, it may he explained 
that of the four successive items composing these numbers, the first 
represents the year, the second the month, and the third the day 
when registration took place, the fourth being the individual 
number of the specimen ; e. (/., means 189S, July, the 
sixth, number five. 

The same symbols are employed as have been customary in tliis 
series of Eritish Museum Catalogues. "[P.] "reads "Presented 
by," " [C.]" equals " Collected by," and "[E.]" signifies "Iteceived 
in Exchange." 

The printing of the Catalogue has taken place, at intervals, 
between March, 19U8, and March, 1912, but the 'Addenda' 
at the end of the volume carry the previously printed sheets up 
to date. The actual date of publication of the volume is March 
23rd, 1912. 

• Aclcnoiuledijments. — My thanks are due, above all, to Mr. Oldtield 
Thomas, •without whose generous support I should not have been 
able to devote practically the whole of my time, for several years, 
to this work, 

I am also under special obligation to the Authorities of the 
United States National Museum, through Mr. Gerrit S. Miller, Jr., 
for the loan without limit of time of a considerable number of 
specimens (including many jtaratypes), chiefly of the genera 


Fleropiis aud Cynopterus, which have been of great help to me 
during ray work. Further, to Dr. P. A. Jeutink, Professor 1*. 
Matschie, and Professor Dr. E. Trouessart for much kind assistance 
during ray visits to the Museums of Leyden, Berlin, and Paris. 

For the loan of type specimens, for information on types or 
other specimens in foreign Museums, or for assistance in any other 
way, I am indebted to Dr. N. Aunandale, Calcutta, Dr. It. Anthony, 
Paris, the Authorities of the Pombay Natural History Society, 
Professor R. Collctt, Christiania, Dr. A. Fritze, Hanover, Professor 
Dr. J. von Kennel, Dorpat, Oberstudienrat Professor Dr. Kurt 
Lampert, Stuttgart, Professor Dr. Einar Lonnberg, Stockholm, 
Dr. Ludwig Ritter Lorenz von Liburnau, Vienna, Dr. Marcus W. 
Lyon, Jr., Washington, Dr. P. Chalmers Mitchell, London, Mr. James 
A. G. Eehn, Philadelphia, Dr. E. lioediger, Frankfort-on-the-Main, 
Senhor A. F. de Seabra, Lisbon, and Yiceinspector Mag. sc. Herluf 
Winge, Copenhagen. The only two illustrations that are not original 
(see above) have been copied by kind permission of, respectively, 
the Council of the Zoological Society, London, and Herr Georg 
lleimer, publisher, Berlin. 

In conclusion I desire to express my thanks to my artist, 
Mr. A. J. Engel Terzi, for the unremitting care and artistic skill 
bestowed by him upon the illustrations. 

K. A. 

British Museum (N.H.). 
January 16tb, 1912. 

.?ij;;:«.;!. i: .i. c ■ 



I. General cliaracters of Megachiroptera ... 

II. Plastic characters of Megachiroptera . 

1. Rostrum, p. xix. 11. 

2. rremaxilhie, p. xx. 

3. Tiifraorbital canal, p. xxi. 12. 

4. Postorhital processes and 
poslorbitnl foramina, 1.3. 
p. xxi. 14. 

.5. Palate, p. xxii. IT). 

G. Tympanic bones, p. xxiii. l(i. 

7. Facial axis, p. xxiii. 17. 

8. Mandible, p. xxiii. 18. 

9. Incisors, p. xxiv. 19. 
10. Canines, p. xxvi. 20. 



Premolars and molars, 
p. xxvii. 

Anomalies in dental 
formula, p. xxxi, 

Palate-ridfres, p. xxxiii. 

Tongue, p. xxxv. 

Wing-structure, p. xxxv. 

Tail, p. xli. 

Calcar, p. xlii. 

Colour of fur, p. xlii. 

Size, p. xliv. 

Secondary sexual cha- 
racters, p. xl V. 

III. Interrelations of the genera of Mcgacliiroptera 

IV. Geographical distribution of Megachiroptera 

1. Distribution of genera, p. Ixv. 

2. Distribution of species and subspecies, p. Isvii. 

3. Remarks on the geographical distribution, p. Ixxv. 

V. Synopsis of the subfamilies and genera of Megachiroptera 
(with a '^Key'' to the genera based on their dental and 
cranial cliaracters only) 





Suboi-derl. MEGACHIltOPTERA. 


Subfamily I. Pteropodinae. 1 

[RorsKTTUS section.] 

1. Eidolon, linjiii 2 

1. diipreanum, Schl. ^- Poll. 7 

2. helvimi, Kn-r 6, 809 

3. siibjeiim, K. And 15 

2. Koliseltus, Gray 1<J 

Roiisettus, Gray ;..... 22 

1 . leachi, A. Smith ' 25, 810 

2. legyptiacus, E. Geoff. . . 29 

3. arahicus, A7id. ir (le Wmt. 33 

4. leschenaulti, I)esm 35, 8l0 

f). seniiiuidus, Eelaart .... 38, 810 

f)bis. ."liortridgei, Thos.. . . . 811 

<), amplexicauclatus, ^. Crcojf. 40 

7. minor, Bobs 43, 811 

«. braohyotis, Bohs 44 

9. celebensis, K. And 46 

Stenonycteris, s»«6<7. «. . 23 

10. kempi, Thos 813 

10 bis. lanosus, Thos 49, 813 

Lissonycteris, suhc/. w. . 23 

11. angolensis, Bocmje 51 

1 1 bis. smithi, Thos 814 

Jl Boneia, Jentink 5o 

1 . bidens, Jentink 58 

4. Pteropus, Brisson 61 

A. Pt. hypomelanus group. i)8 

1. hypomelanus, 7"f)/iw(. .. 101 
ft. f^eminovum, vT/tV/^'r .. 106 

I. enganus, MiVer '. 107 

<■. condorensis, iV^ 110 

d. canus, K. And. ...... 1 1 ^ 

«. lepidus. Miller 115, 815 

f. mmectens, K.. And. .. 116 

y. robinsoni, K. And. . . 815 

(/. tomesi, Pet. 119, 816 

/i. cagayanus, Mearns . 121 

u macassaricus, Ileude . . 124 

). hyTpom6\ar\\\!', Temin. . . 127 

'k. luteus, 7t. .4W 128 

2. spt^ciosus, K. And 132 

S. mimu?, 7v'. vlnf/ 133 

4. pallidus, Tcmni l^O 


5. griseus, E. Genjf. 137, 816 

5 bis. pumilns. Miller .... 816 

6. satyrus, K. And 142 

7. faiinulus, Miller 14.3 

8. admiralitatum, 'J'hos. .. 144 

9. colon us, K. And 147 

10. solomonis, Thos 148 

11. brunneus, Bvbs 149 

12. oraatus, Gray 153 

13. auratus, K. And 156 

14. dasymallus, Temm lo9 

lo. formosus, /', L. Sclater.. 163 

15 bis. Hops, Thos 817 

16. subniger, Ken 164 

B. Pt. niariannus group .. 172 

17. pelewensis, K. And 173 

18. yapensis, A. And 174 

19. ualanus. Pet 177 

20- mariannus, Besm 178 

21. loochoensis, Gray ...... 181 

22. vanikorensis, Q. ^ G. . . 184 

23. tonganus, Q. ■^- G 186 

24. geddiei, Maeyill 189 

C. Pt. caniceps group .... 192 

25. dobsoni. A'. .47jrf. 192 

26. caniceps. Gray 194 

27. arcentatus, CrVoy 197 

D. Pt. ruf us group ...... 200 

28. nifus, E. Geoff. 202 

a. rufus, E. Geoff. 204 

b. priuceps, K. And. .... 208 

29. comorensis, Nic(dl .... 208 
29 bi*i. voeltzkowi, Af«/«c//?« . 818 

30. sevchenensis,71/?7wp- ii(/H'. 212 

31. ftldabrensis, True 213 

32. niger, Kerr 215 

E. Pt. inelanotus group . . 223 

33. melaiJotus, Blyth 224 

34. tvtleri. Mason 227. 820 

35. niadicus, Mitler 229 

36. modigliaiiii, Thos 232 

37. natalis, Thos 233 

F. Pt. melnnopogon group . 237 

38. melanopogon, Pd 238 

.39. aruensis. Pet 241 

40. keyensis, Pet 246, 821 

41. livingstonei, Gray 247 

(i. I't. rayneri group .... 2oO 

42. cognatus, A'. And 251 



43. rayneri, Grnj/ S-'iS 

44. rubianus, K. Awl :?oo 

45. IftVfllaiius, K. And 258 

4(). graiidis, T/io.t 259 

47. chrysopvootu.s, Temm. . . 2(J0 
H. I't. loinbocen«is group . 2(i5 

48. lombocensi.i, Dob/i 2Gt> 

49. solitarius, K. And., 269 

50. rodricensis, Dohs 273 

51. molossinu.", Temvi. .... 275 
I. Pt. samoensis group . . 280 

52. nawaien.sis, frrai/ 280 

5-3. samoensis, Peale 284 

54. anetianus, G?yn; 288 

J. Pt. pselaphon group . . 293 

•55. insularis, Homhr. 4" J«cq. 295 

56. phreoceplialus, Thos 298 

57. pselaphon, Lay 301 

58. pilosus, K. And 306 

59. tuberculatum, Pet 309 

60. leucopteru.s, Temm 311 

K. Pt. temmincki group . . 315 

61. temmincki, Pet 316, 822 

62. capistratus. Pet 319 

63. personatus, Temm 321 

L. I't. vampyrus group . . 324 

64. piganteus, Briinn 326 

a. gigauteus, Bribm 329 

b. leucocephalus, Hodys. . 333 
6.5, ariel, G. M. Allen 335 

66. lylei, K. And 339 

67. interniedius, K. And. . . 340 

68. vampynis, L 343 

fl. maiaccensis, A'. And. . 346 

b. vampyrus, L 349 

c. pluton, Temm 353 

d. edulis, E. Geoff. 356 

e. nataiife, K. And 358 

f. ItinensLs, Mearns .... 359 
M. I't. alecto group 3G3 

69. aterrimus, Matxchie .... 363, 822 

70. alecto, Temm 365 

71. morio, K. And. 370 

72. gouldi. Pet 370 

N. Pt.couspicillatus group. 375 

73. chrysauchen, Pet 375 

74. conspiciilatu8, Gould. . . . 378, 823 

75. ocularis. Pet 381, 823 

O. Pt. neoliiberiiicus group 384 

76. papuanus, Pet. ^- Jioiia . 38-j, 823 

77. neohibernicus. Pet 387 

P. Pt. macrotis group .... 392 

78. epulnriiis, Panisai/ 392 

79. m.acroris, Pet. . . . " 396 

80. poliocephalus, Temm. . . 397 
Q. Pt. scapulatus group . . 402 

81. scapulatus, Pet 403, 824 

82. woodfoiHli, Thos 407 


5. Acerodon, Jour dan 412 

1. celebensis, Pet 417 

2. mackloti, T«>!?)i 418 

n. mackloti, Temm 419 

h. Horesii, Grnii 420 

c. alorensis, K. And 423 

3. gilvus. A'. And. 423 

4. humilis, K. And 424 

5. jiibatus, BseJiRch 426 

a. jubatus, Eschsch 427, 824 

h. miudanensi.s, K. And. . 429 

6. lucifer, Elliot 432 

6. Pteralopex, Thos 432 

1. ancepf, K. And 437 

2. atrata, Thos 439 

7. Styloctenium, Mntschie . . 442 
1. wallacei, Grai/ 44-5 

8. Dobsonia, Palmer 448 

1. minor, Dobs 460, 824 

2. exoleta, K. And 461 

3. pannietensis, T)e Vis .... 463 

4. moluccensis, Q. S,- G 464 

5. magna, Thos 466, 825 

6. pevoni, A^. Geoff. 467 

7. surabana, K. And 471 

8. vii-idis, Heude 471 , 825 

n. umbrosa, Titos 825 

h. viridis, Heude 826 

9. crenulata, K. And 47-3 

10. praedatri.-?, A'. And 474 

11. inermis, K. And. 475 

12. nesea, K. And. 476 

[Epomophokus section.] 

9. Plerotes, K. And 483 

1. ancbietSR, Senbra 486, 837 

Epomop.s, Gray 487 

franqueti, Tomes 494 

a. strepitans, A'. And. . . 496 

h. franqueti, TodiC^ .... 497 

bupttikoferi, Matschie . . 499 

dobsoni, Bocage 500 

IIyp*ignatluis, H. Alien . . 501 

, monstrosus, H. Allen . . 506, 827 

Epomophoru.", />^'7f?!(-/< .. 514 

wahlbergi, Sund. ...... 521 

a. haldomani, Hidl.owell . 522, 827 

b. wahlbergi, Snnd 526 

labiatus, Temm 529 

minor, Dohs. .......... 531 

anuru.a, HeugHn. ....... 532 

crypturus. Pet 535, 827 

g.ambianua, 0//('Wy ^^^'^ ' 

7. angolensis, Gray 542 

8. pou.'>arguesi, rz-ot/Ms. .. 543 
13. Micropteropus, Matschie . 654 

1. pusillus, Pet. 557 







14. Nanonycteris, Matschie . . 
1. veldkanipi, jenttnic .... 

15. Scotonycteris, Matschie . . 
1. zenkeri, M<itschie 

16. Casinycteris, Thos 

1. argynnis, Thus 

[Cynopterus section." 

17. Myonj'cteris, Matschie . . 
Myoiiycteris, Matschie . 

wroughtoni, K. Atid. . . 

leptodon, A'. And 

torqnata, Dobs 

Phy!?c-ti8, siibff. n 

brachycephala, Bocuye . . 

18. Cynopterus, F. Cuv 

sphinx, Vahl 

a. sphinx, Vahl 

h. gancreticus, K. And. . . 
c, titthsecheilus, Temm. . . 

2. brachyotis, S. Miiller. . . . 

a. angulatus, 3Iillcr .... 

b. brachyotis, S. Miiller. . 

c. javamcus. A'. And. . . 

d. insularuni, K. And. . . 
■ e. ceylonensis, Grai/ .... 

■ ' - f. niinUtus, Miller 

//. bracbysoma, I>ol>s. . . 
h. scherzeri, Zelebor .... 

3. major. Miller 

4. borsfieldi, Grai/ 

a. liorsfieldi, Gray .... 
h. lyoni, nom. n. 

5. harpax, Thos 

0. princeps. Miller 

19. Ptenochirus, Pet.. ..... 

1. jagori, Pet 

20. Megferops, Pet 

1. ecaudata, Temm 

21. Dyacopterus, yew. M. .. 
1. spadiceus, Thos., , , , . . 

22. Balionycteris, Matschie 
1. maculata, Thos 

2;3. Chironax, gen. n 

1. melanocephalus, 2'emm. 

24. Thoopteriis, Matschie . . 
1. nigrescens. Gray .... 

25. Penthetor, gen. n 

1. lucasi, Dobs. 

20. Sphserias, Miller 

1. blanfordi, Thos 

27. Nyctimene, Borkhaiisen 

1. papuanua, K. And. . . 

2. albiveutei', Gray .... 





632, 827 


3, minutus, A'. And 701 

4. variiis, K. And 702 

6. cyclotis, K. And 703 

5 bis. certans. A', And 828 

6. cephalotes. Pall 703 

7. geiuinus. A'. And 709 

8. major. Dobs 710 

9. scitiilus, K. And 711 

10. lullula;, Thos 713 

] 1. robiusoni, Thos 714 

12. aello, Thos 715 

Subfamily II. Macroglossinae. 723 

[EoNYCTERis section.] 

28. Eonycteris, Dols 728 

1. spelaea, Dobs 734 

2. major, K. A7id. , 730 

3. rosenbergi, Je?itink .... 737 

29. Megaloglossus, Payenstecher 738 
1. woevrnMim, Payenstecher. 742 

30. Macroglossus, F. Cuv. . . 746 
1. minimus, E. Geoff. .... 755 

a. minimus, E. Geoff. . . 757 

b. sobrinus, A'. And 760 

2 Iagochilu8, Matschie .... 7(52 

a. lagochilus, Matschie . . 703 

b. nanus, Matschie 765 

c. pygmaius. A'. And. . . 707 

d. microtus, K. And 767 

31. Sjconycteris, Matschie .. 771 

1. crassa, Thos 775 

a. papuana, Matschie .... 777 

b. keyensis, A'. And. .... 779 

c. linschi, Matschie .... 779 

d. crassa, Thos. 780 

e. major, K. And 780 

2. australis, Pet 781 

3. naias, A'. And 785 

[NoTOPTERis section.] 

32. Melonycteris, Dobs 785 

1. melauops, Dobs 789 

33. Nesonycteris, Thos 790 

1. woodfordi, Thos 792 

34. Notopteris, Gray 793 

1. macdonaldi. Gray 797 

2. neocaledonica, Trouess.. . 709 

Subfamily III. Harpyio- 

nycterinae 799 

35. Ilarpyionycteris, Thos. . , 799 
1. whiteheadi, Thos 805 

List of Illustrations 835 

Alphabetical Index = 839 


I. General ckakacters of Mkgacjiiroptera. 

The prototype of tlio Chiroptera must have possessed all tlio 

mo.^t primitive features preserved by any living or extinct form of 

!Me.gachiroptera and Microchiroptera. Among its more important 

characters, therefore, must have been these : — 

Skull. — In general shape probably essentially like that of a 

liouscttus or Pteropiis, with the following differences : the bony 

palate was not produced backward behind the tooth-rows (compare 

Insectivora and Jlicrochiroptera generally) ; the infraorbital canal 

long {cf. the Notopteris section of Megachiroptera, and somo 

Microchiroptera); the postorbital processes undeveloped (as in the 

majority of Microchiroptera"); tlie facial portion of the skull not, 

or only inconspicuously, deflected against the basicranial axis (as 

in nearly all genera of the Epomopliorus and Crjnopiei'us sections 

of Megachiroptera). 

n .-.• T^ ,-^e ^ t,ui ^' i" — ° P' — p^ p' m' m= nv\ 
Ventition. — Dental tormula probably .-^-^ *, 

I1I2I3CP1— P3P4m, ^2^3 

this being the completest formula found in any bat (i' may have 
been ])resent, though lost in all known species) ; molar structure 
typically " Insectivorous," as in Microchiroptera with unmodified 

Fore-limbs and memhravcs. — Tuheratlum majus and miiuis of 
humerus relativel}' small, the former not articulating with scapula 
(cf. Megachiroptera and some primitive Microchiroptera) ; deltoid 
crest of humerus weak (cf. Megachiroptera). Ulna not more 
reduced than in Megachiroptera. Articular surfaces of the bones 
of the hand (earpo-mstacarpal, metacarpo-phalangeal, and inter- 
phalangeal joints) not more modified than in Megachiroptera ; 
trapezium large (cf. Megachiroptera) ; second digit rclalively 
independent of third, with three phalanges and a well-developed 
claw (cf. Archcmpteropus f and typical Pteropodida). Wing-i 
membranes from tlaiiks and inserted i)osteriorly on first toe. ' 

Tail. — Long and embedded in interfemoral (cf. ArcTiceopteropus, 

* On the liouiologies of the iiiissing premolars in Chiroptera, see Oldfielcl 
Thouias. Ann. & Maj;. K. II. (8) i. p. 3i'j (1908). On the missing upper 
incisor, K. Andersen, P. Z. S. 1908, p. 205. 

t On Aixhieopteropua traiisieus (Upper Oli^ocene, Monleviule, Italy) see 
IMesfhinelli, Atti R. 1st. Veneto, Ixii. p. 1329, pi. is. (1903). Note on il3 
wine-struolm-e, infra, p. ssxvii. 



iVotop<<?Hs, typical Microehiroptera), number of free caudal vertebrae 
probably at least ten *. 

Tbe characters in wbich living Megachiroptera differ from the 
above scheme of the ancestral form may be divided into two classes, 
those that have become j^.recZ, i.e. firmly and invariably established 
in all living genera, and those that are plastic, i. e. subject to 
variation from genus to genus. 

The modifications common to all living members of the Suborder 
are these : — 

(1) The bony palate is elongated behind the tooth-rows and the 
zygomatic process of the maxillary bone (the practically complete 
absence of the postdental portion of the palate in a single genus, 
Casinyrteris, is without doubt a secondary modification, Casinycteris 
being closely related to a form (S colony cter is) with normal Mega- 
chiropteran palate) : 

(2) Postorbital processes are developed : 

(3) The molar structure is modified from the common Insecti- 
vorous type into the ordinary Megachiropteran type as follows 

i~^ 4*5 

Fig. I. — Typical rnolar structure of Megachiroptera compared with 

that of Insectivora. 

A. right upper m^, B. left lower m^ oC Talpa eiiropma (B.M. 

A', right upper m', B'. left lower mi o{ Rauseitus (egyptiacus (B.M. 

A and B f , A' and B' f. ' 

For explauation of nuiiibering of cusps see footnote t below. 

(fig. I.): in the upper molariform teeth cusps 1, 2, and 3t have 
disappeared, cusp 4 and the reduced cusp 5 form together a longi- 
tudinal ridge along the outer side of the teeth, cusps 6 and 7 

* Most of the characters enumerated above have already been suggested by 
Herluf Winge in his " Jordfundne og nulevende Flagermus {Chiroptera) fra 
Lagoa Santa, Minas Gei-aes, Brasilien ; med Udsigt over Flagermusenes 
indbyrdes Slsegtskab" (E Museo Lundii, vol. ii. pt. i. p. 27; 1892). For a highly 
instructive account of the modifications that have taken place in the develop- 
ment of the Chiropteran type from some primitive form of Insectivora, see 
pp. 18-23 of Winge's memoir. 

t The molar cusps are named in accordance with Herluf Winge's theory 
(" Om Pattedyrenes Tandskifte, isfer med Hensyn til Tjendernes Former," 
Vidensk. Meddel. naturhist. Foren. Kj0benhavn, for 1882, pp. 16-18, pi. iii.). 
The three cusps, labial in the upper but lingual in the lower teeth, that form 
the tips of tbe W of a typical Insectivorous molar are termed, in antero- 
posterior direction, respectively 1, 2, and 3, cusp 2 being the oldest, homologous 
with the single cusp of a Reptilian tooth; the two cusps forming the bases of 
the W are named 4 and 5 ; and the "' heel " of the upper molars, when single, 
cusp 6, when double, cusps 6 and 7. 



a similar ridgo along the iuner side, the two ridges being separated 
by a median longitudinal depression ; in the sirailarlj* shaped lo'vrer 
niola.riform teeth the inner ridge is formed by cusps 1, 2, and 3, 
tlie outer by cusps 4 and 5 : 

(4) ig and m' are lost : 

(5) The second and, particularly, the third phalanx of the 
second finger are somewhat reduced in length. 

The principal plastic characters of the Megachiroptera are 
discussed in the next paragraph. 

II. Plastic characters of Megachiroptera. 
1. Rostrum. 

Varies considerably in length, much less so in sliape. 
Length. — If measured from the front of the orbit to the extremitj- 
of the nasal bones, the cranial rostrum is : — 

(1) Longest, i. e. from about f to nearly ^ of the skull, in 
Hi/psir/nathits and some species of Ejiomophorus, in both genera 
relatively longer in males than in females ; the male ffypsignafhus 
is the longest-faced of all Fruit- bats : 

(2) Medium or somewhat shortened, i. e. from about g to |^ of 
the skull, in : — (a) the genera of the Bousettus section ; (b) all 
Macroglossince ; (c) within the EpomopiTiorus section in the genera 
Plerotes and Epomops, and in some (the smaller) species of Epomo- 
phorus; (d) within the C'ynopferus section only in Myonycteris 
and Spharias : 

(3) Shortest, i. e. about I— I of the skull, in :— («) the following 
four genera of the Epomoplwrtis section, Micro pteropus, Nano- 
nycteri.'; (rostrum rather more than ^ of skull), Scotonycteris, and 
Casinycieris : (6) all genera of the Cynopterus group, except 
Myonycteris and SpTK^rias. 

Briefly summed up : a moderately long rostrum is characteristic 
of all Eonsettine and Macroglossine Fruit-bats, a very short of 
nearly all Cynopterine genera, while the Epomophorine is the only 
section showing any degree of variation in the length of the 
rostnira, from that of Plerotes (moderate), on the one hand to that 
of Ilypsrgnathus (longest known), on the other to that of Casi- 
nycieris and allied genera (shortest). 

No Fruit-bat with a very short rostrum (third stage above) has 
the full Megachiropteran number of cheek-teeth (the formula being 
eilher j, j, or A\ but this rule cannot be reversed, that is, a 
reduction in the number of teeth is by no means always, not even 
usually, associated with a shortening of the rostrum. Fruit-bats 
with a moderately long or very long rostrum (second and first 
stage above) exhibiting any variation of the dental formula from 
the highest number of cheek-teeth (|) to the lowest (0. 

Shape.— As a general rule the rostrum is conspicuously tapering 



jinteriorly, and tlio aberrations from this typical (Insectivorous- 
like) sha[)e are, apart from Ilypsignatlms, but few and not very 
peculiar. The rostrum is unusually deep (truncate) in front in 
JSIeqarops and Nyctimene (both of the Cynoptevus section) ; in 
Niictimene the medial portion of the nasals is produced forward 
and downward as a triangular projection firmly united with the 
anterior edge of the mesothmoid cartilage (evidently acting as a 
support of the nasal tubes; c/. fig. 61, p. 684). In Fruit-bats 
with weak dentition (narrow cheek-teeth) the rostrum is lower and 
thinner than usual ; compare Stenonycteris (fig. 3, p. 49) with 
llousettus (fig. 2, p. 17), Pteropus scapulatas (fig. 18, p. 404) with 
the average Pteropus skull, Sphcprias (fig. 60, p. 672) with the 
ordinary Cyaopterine type of skull (fig. 48, p. 588), and all Macro- 
(/lossince, except the more heavily-toothed Eonycleris. The only 
strikingly peculiar modification is seen in males of Hypsignathus 
{fig. 33, p. 502) 4 the rostrum is greatly increased in size and 
jjarticularl}' in depth, its dorsal and ventral profiles subparallel, its 
lateral surfaces concave (a large subcutaneous air-sac is present on 
either side of the rostrum), and a strong vertical crest runs from 
the nasals to the alveolus of the canine (supporting a cutaneous fold 
of the upper lip) ; in females of the same genus the rostrum is 
somewhat similarly, but much less excessively, modified. 

2. Premaxillce. 

Vary in shape (breadth, degree of reduction), in direction, and 
in the mode of interconnection of the alveolar branches. The 
palatal branches of the premaxilliB are always absent. 

Breadth. — The three Macroglossine genera Mdonycteris, Ntso- 
nycteris, and Notopteris exhibit what is probably the most primitive 
form of the premaxilke : the ascending branch is unreduced in 
breadth at its upper extremity, gradually narrowing interiorly, so 
that the breadth of the bone near the alveolar margin is only 
one-half or one-third of its breadth at the upper extremity (fig. 76, 
p. 791). The next stage is shown by genera in which the ascending 
branch is considerably reduced in breadth above, being quite or 
nearly as narrow above as near the alveolar margin (ex. Pteropus 
and allied genera ; Epumophorus ; Eonycttris ; Meijaloylossiis ; 
Macroqlossus; &c.) ; a still higher degree of reduction by genera 
in which the bone is tapering to a point above, its upper extremity 
often slightly curved forward (ex. nearly all geneia of the Cyno- 
pterus section) ; and finally, the whole of the ascending branch may 
become so thin as to be almost linear throughout (Dohsonia). The 
whole of the premaxillary region is increased in breadth in Bypsl- 
(jnathus (owing to secondary modifications in the shape of tlie 
rostrum). In Nyctimene (fig. 61, p. 684) the alveolar branches 
are unusuall}' deep (vertically), the ascending branches so short as 
not to reach the nasal bones (extremity of rostrum modified, for 
support of the peculiar nasal tubes in this genus). 

Direction, — As a rule the ascending branches of the prcmaxillse are 



ri carl}- vertical oroulyslightlvproclivoiis. A somewhat, or even much, 
liighcr degree of proclivity is seen in Fhrotes, Epomops, SiJucrias, 
all Macrofjlossiiw; except Eomjcteris, and in Harpyioivjctens. 

Interconnection of branches. — In the large majority of Fruit-bats 
the right and left alveolar branch of the preraaxillte are in simple 
contact anteriorly, the suture between them being permanent. 
In certain genera, however, they become, at an early age of tho 
individuals, united by synostosis, i. e. firmly and solidly ankylosed 
together, without any trace in the adult (and, as a rule, not even 
in the semiadult) of an inter-premaxillary suture ; this is the case in 
Lissonycteris (subgenus of liousettus), Fteralopex, and Harpyio- 
ni/cteris, all of the Bouseitus section, in Hy psiynathus among the 
lipomophorine Fruit-bats, in Dyacopterus, C'hironn.v, and Ni/ctimene, 
of the Cynopterus section, and in AJeyaloyhssus, 2Iaeroylossus, 
Syconycteris, and Xotopteris, among the JIacroylossinee. The 
opposite extreme is represented by a few genera in which the 
premaxilloe are not even in contact anteriorly, but distinctly spaced, 
connected by fibrous tissue only (constantly 'in Eidolon and Boneia ; 
occasionally in Eonycteris and Mdonycteris). 

3. Infraorbital canal. 

In Insectivora, as well as in primitive Mammalia genenilly, the 
infraorbital canal is long, its anterior aperture, the infraorbital 
foramen, situated a considerable distance in front of the orbital 
cavity. The only forms of Megachiroptera which in the length of 
the infraorbital canal approach to this primitive condition are the 
three genera Melonycteris (fig#75, p. 786), Nesonycteris (fig. 76, 
p. 7U1), and Notopteris (tig. 77, p. 795), all closely interrelated and 
belonging to the subfamily JIacroylossince. In all other Fruit-bats 
the canal is considerably shortened, its outer wall, as a rule, a 
narrow, often sublinear, bridge of bone, and the foramen situated 
nearly vertically below or only slightly in front of the orbital 

4. Posiorbital processes and postorbilal foramina. 

Processes. — The postorbital processes are generally small or 
moderately strong, and if so there is scarcely any trace of corre- 
sponding lower processes from the zygoma. In a small number of 
forms, all with heavy dentition (chiefly species of Ftcropus and 
allied genera), they are stronger than usu'al, reacliing about halfway 
between frontal and zygoma, and the lower processes are more or 
less conspicuously developed. Earely (a few large species of 
Pteropus and Acerodon ; Pteralopcx) they are, at in aged 
individuals, so long as to join the lower processes and form a 
complete ring round the orbit. 

Foramina. — The base of the postorbital process is nearly always 
pierced by a relatively large foramen (no doubt homologous 
with the supraorbital notch or foramen of the human skull, for 


Nerviis and Vasa supraorhitalia). The oulj section of genera 
in which exceptions to this rule occur is the Cynopterus section ; 
the postorbital foramina are, within that group of Fruit-bats, 
normall}" developed only in Myonycteris, Cynopterus, Ptenochirus, 
and Aleycerops, minute (tending to disappear) in Dyacopterus, 
generally absent (or, it' present, minute or traceable only on one 
side of the skull) in Nyctiinene, always absent in Balionycteris, 
Chironax, Thoopterus, Penthetor, and Sphcerias. 

5. Palate. 

Pteropus (fig. 6, p. 62) is probably one of the genera that shows 
the Megachiropteran palate in its least modified form. The inter- 
dental palate is only moderately broad, gradually narrowing in 
front (the tooth-rows distinctly converging anteriorly); the post- 
dental palate about one-fourth of the total length of the palate, its 
lateral mai-gius forming almost straight lines converging posteriori)-. 
In Ho usetius and Boneia the interdental palate is relatively broader; 
an even greater increase of breadth is seen in Epomops, and in 
Plerotes this portion of the palate reaches its maximum of breadth 
(bearing a curious resemblance, in general outline, to that of the 
Carnivorous genus Proteles !). An opposite course of development 
is taken by Epomophorus, in which the palate is unusually narrow. 
If the lower canines are slanted outward, this necessitates a greater 
breadth of the palate anteriorly, between the upper canines, and 
therefore more nearly parallel tooth-rows (ex. Boneia, Uypsi- 
gnatJius, Macroglossus). 

The postdental palate is in the majority of genera essentially 
similar to that of Pteropus described above ; it may be broader or 
narrower, shorter or longer, more rapidly narrowing posteriorly 
or more nearly parallel-margined ; in Nyctiraene it is somewhat 
pandurate in outline, i. e. distinctly constricted at middle (owing 
to the unusually broad mesopterygoid fossa ; compare the presence 
of nasal tubes in this genus)- But these variations are com- 
paratively trivial as compared with the modifications exhibited by 
some genera of the Epomophorus section : — In Plerotes, Epomops, 
Hypsigiiathus, and Scotonycteris it is relatively simple in shape, 
though in the two former genera rather broader than usual ; but 
in Epomopliorxis (fig. 36, p. 515) it is deeply depressed posteriorly, 
and its posterior free margin high and prominent ; m Micro pteropus 
(tig. 38, p. 555) it is abruptly narrowed behind the roots of the 
zygomatic processes, the posterior depression is shallower, the free 
margin prominent ; in Nanonycteris (fig. 40, p. 560) it is short, 
but unusually broad, longitudinally depressed on either side, 
slightly convex between, and flattened at the extreme posterior 
extremity, the free edge not prominent ; and, finally, in Casinycteris 
(fig. 43, p. 569), a genus in other respects closely similar to Scoto- 
nycteris, the postdental palate has practically disappeared, the 
mesopterygoid fossa extending forward very nearly to the level of 
the posterior molar, a modification unique in Megachiroptera and 
more closely recalling the type of palate found in Microchiroptera, 


6. Tympanic bodies. 
Alwajs annular, but the ring varying a little In breadth (rather 
broader than usual in liousettus, Dobsonia, Eouyctey-is). The only 
peculiar modification is the development of a short bony auditory 
meatus in Eidolon (fig. 1, p. 3 ; cf. Miller, 'Families and Genera 
of Bats,' p. 55, fig. 7 B), a genus closely related to liousettus. 

7. Facial axis. 

Normally the facial axis if projected backward is nearly parallel 
witli the basicrauial axis, i. e. the former forms an angle with the 
latter of not much less than 180 degrees, and the line of the alveolar 
margin of the upper tooth-row if continued backward passes through 
the lower edge of the occipital condyle or even somewhat below the 
condyle ; such is the case, with rare exceptions, in all genera of the 
Epomophorus and Cynopterus sections (ex. fig. 36, p. 515, and tig. 48, 
p. 588). In the Eousetto-Pteropine section generally the face is 
more distinctly, sometimes even very conspicuously detiected on 
the basicrauial axis, so that when the rostrum is kept horizontal 
the axis of the brain-case points obliquely backward-and-downward, 
and the posterior projection of the alveolar line passes through the 
middle or upper margin of the occipital condyle (ex. fig. 1, p. 3 ; 
fig. 6, p. 62) or even through the middle of the supraoccipital 
(fig. 3, p. 49 ; fig. 18, p. 404). The maximum of facial deflection 
is seen in the majority of Macroglossincn, in which the face is bent 
downward to such degree that the alveolar line if continued pos- 
teriorly would pass through the brain-case considerably above the 
lambdoid crest (fig. 70, j). 748). 

As a general rule the deflection of the face is greatest in genera 
or species with weak dentition ; thus it is considerably greater in 
the narrow-toothed Stenonycteris (fig. 3, p. 49) than in typical 
Rousettus (fig. 2, p. 17); in Pteropus the deflection is always 
noticeable, but least so in the heavy-toothed species (ex. fig. 15, 
p. 302, Ft. p)selaphon), and unusually great in the small- and 
narrow-toothed Ft. scapulatus (fig. 18, p. 404) and Ft. woodfordi 
(fig. 19, p. 408) ; the only Epomophorine genus with conspicuously 
deflected facial axis is the very weak-toothed Flerotes (fig. 28, p. 484), 
and the only Cynopterine genus with similarly deflected face is the 
weak-toothed 8p>ha;)-ias (fig. 60, p. 672) ; within the subfamily 
MacrocjlossincB the deflection is smallest in the relatively strong- 
toothed Eonycleris (fig. 66, p. 730), greater in Meyalaglossus (fig. 68, 
p. 739), which in the degeneration of the dentition is transitional 
between Eonycleris and Macroglossus, very great in all other 
genera, all of which are characterized by unusually narrow or 
small cheek-teeth. 

8. Mandible. 
The heavier the dentition, the broader and deeper and more 
vertically ascending is the coronoid process of the mandible, the 
stronger its angular process, and the higher above the alveolar line 
its condyle ; the weaker the dentition, the narrower and lower and 
more backwardly sloping is the coronoid process, the feebler the 


angular process, and the lower dowu the position of the condyle, 
Tliere is within the suborder every gradation from the unusually 
heavy mandible with nearly vertical coronoid process, deep and 
broadly rounded angular jjrocess, and the cond\le situated high 
above the alveolar line, of Pteratopex (fig. 21, p. 433j, Acerodon 
(fig. 20, p. 413), the heavy-toothed species of Pteropus (fig. 14, 
p. 281) ; fig. 15, p. 302), Dohsonia (fig. 24, p. 449), and most genera 
of the Cyuopterine section (fig. 48, p. 588 ; fig. 61, p. 684), through 
the intermediate conditions as exhibited in the Epouiophorinc 
section (fig. 30, p. 488 ; fig. 40, p. 560), to the mandible as shaped 
in the narrowest-toothed Macroi/Iossina; (fig. 70, p. 748), with 
its short, thin, and posteriorly directed coronoid process, weak 
angular process, and condyle situated considerably below the level 
of the alveolar line. 

The symphysis of the mandible is in the Pteropodince usually 
obliquely ascending, more rarely (ex. Dohsonia, Nyctimcne) sub- 
vertical ; in the Macroglossince as a rule more nearly horizontal. 
In some genera of Macro(jlossince {Eonijdcris, Mei/alor/Jossus, Macro- 
(jlossus, Mdonycteris, Nesomjcteris) there is a more or less clearly 
pronounced tendency to a development of a longitudinal keel 
along the antero-inferior surface of the symphysis. 

9. Incisors. 

Incisor formida'. — i^ is lost in all Chiroptcra ; ij is present in 
many Microchiroptera, but lost in all Mcgachiroptora (the incisors 
of Arclia'opteropus are unknown) ; no Fruit-bat, therefore, has more 
than % — 5 incisors, but some have less. The following five incisor 
formuliB occur : — 

i- iMM 2 2 

.-. = , the normal formula, found in all genera except 

'^ 'i 'i '-^ ^ , ^ 

those enumerated below. 

j2 i^ 1 1 . 

= , in one genus, Boneia (allied to lioitsetfus). 

\h\h ^' ^ 

42 \\ p V- 9 2 . ■ 

= , in one genus of the Ptcrojms subsection, viz. 

Sh/locfe7iium ; in four genera of the Cynopterus section, 

ptcnochirus, Megcurops, Balionyctcris, and Penthetor; 

and in two Macroglossine genera, I\esonycicris and 

JS'otoptcris (in the latter i^ is deciduous). 

;2 i^ 1 1 

= , in Dohsonia and Ilarpyionyctens^ both aberrant 

i., ij 1 ■'^ 

genera of the Pousethis section (on the homologies of 

the missing incisors in these two genera, see footnotes 

p. 452 and p. 801). 

- i'i' - 1 1 

Z = , in Kiictimcnc onlv, a i)eculiar]v modified genus of 

the Cynopterus section. 


In Pkropus i, is always conspicuouslj^ smaller than i„, and in 
some species {I't. lomhocensis group) so luuch reduced as to be 
almost rudimentary ; this foreshadows Stylodenium, in which ij is 
completely lost, and Dobsonia and Ilui-pyionijcteris, in which both 
ij and i' are lost. In those genera of the CynojJients section which 
have the normal number of incisors i, is Aveaker than i„, and i" 
iuintly shorter than i' ; this leads to those genera of the same 
bcclion, viz. I'tenochirus, Meijcerops, Balionijcteris, and I'eniheior, in 
which ij has disappeared and i" is shortened to about half the length 
of i' ; and to JVyctimene (lower canines moved forward to extremity 
of jaw), in which all lower incisors as well as i" have disappeared. 
In the Macroglossine genus MeJonyctens \ is smaller than i.^, in the 
closely related J^esomjcteris and Is'otoptivis it has disappeared. 

Differentiation. — In the majority of Fruit-bats the incisors are 
small, thin, terete (styliform) or subterete, the crown only slightly 
or even indistinctly differentiated from the shaft, the cutting-edge 
of the lower incisors often faintly bilobed. Normally the incisors 
above and below are placed approximately vertically, those in the 
upper jaA\" in contact with each other or narroAvly spaced, but with 
a relatively wide diastema on either side between the incisors and 
canines, those in the lower jaAV crowded and in contact with the 
canines. The principal variations, to be described below, from this 
predominant type of incisors are, briefly summed up, these : (1) the 
crown of some or all of the incisors may become peculiarly diifer- 
entiated in shape, (2) conspicuous posterior ledges may be developed, 
or (y) some of the incisors may be enlarged or reduced; further, 
(4) the direction of the incisors may be altered, and (5) the spacing 
of the incisors may be conspicuously unequal. 

In Fteroinis (and the closely allied Acerodon) the upper incisors 
are larger than usual, the crown distinctly differentiated, posterior 
basal ledges conspicuously developed, sometimes so much so as 
to form a noticeable shelf (7^<. pscZo^^/ion group), i, always larger 
than ij, either owing chiefly to a reduction of ij, which may 
become nearly rudimentary (Pt. loinhocensis group), or chiefly to 
an increase of i^ {Ft. samoensis and iisda^ihon groups). Some of 
these tendencies are further developed in the related 
I'teralopex : the posterior ledge of the upper incisors is very large, 
shelf-like, rendering the antcro-posterior equal to or greater than 
the transverse diameter of the teeth, and i, is from twelve to fifteen 
times the bulk of i^, its cutting-edge unequally trifid (middle cusp 
much the broadest). 

In Splurrias {Cynopterus section) both the upper and lower in- 
cisors arc triangularly pointed. In Syconycteris {Macroijlossivcp) the 
upper incisors are larger than usual, the crown well-differentiated, 
narrowly chisel-shaped, i„ considerably higher and broader than i^, 
with obliquely triangular crown. 

In those genera (Dolsonict, Ea)pyionyci^ri.i, ]\''yctimenc) which 
have only [ — j or ^ — ^ incisors, and in which the lower canines are 
situated close together or in actual contact at the anterior extremity 
of the mandible, the single pair ol upper incisors is acted upon by 


the tips of the lower caiiines and somewhat altered in shape, the 
crown being more or less obliquely bilobed. 

The normal vertical direction of the incisors is changed into a 
strong proclirity in SjjfKerias, Syconycteris, and Harpyionyctens. 

Aberrations from the normal crowded or equidistant arrangement 
of the incisors are seen in several genera of Maeroglossince : in 
Melonycteris i^— i^, in XesonycUris and Xoiopteris \^—i\ and in 
Macroglossus both i^— i' and i,— i^ are unusually widely spaced. 

In Hypsignathm {Epomophorm section) the lower incisors do not 
bite against but close some distance in front of the upper. 

Deciduous incisors.— \' (rudimentary, barely piercing gum) is 
deciduous in XotopteAs ; i^ (relatively well developed) otten 
deciduous in Epomops (for details see p. 489, footnote) ; i^ (the 
single lower incisor, almost functionless) sometimes deciduous 
in Dobsonia {D. viridis and crenulata), perhaps so in Harpyio- 

10. Canines. 

Apart from minor variations in length and bulk, the canines may 
be moditied : (1) by an enlargement of the cingulum, (2) by the 
development of secondary cusps, or (3) by the development of 
longitudinal grooves on the crown of the upper canines ; further, 
(4)°the direction of the canines may be changed (proclivity ;_ out- 
ward or outward-and-backward slant of lower canines), and (5) the 
position of the lower canines may be changed. 

Ciru/ulum.—As a general rule, in Fruit-bats with weak dentition 
the cingulum of the canines is thin or obsolescent ; on the other 
hand, the heavier the dentition, the thicker and more prominent 
the cingulum. In Pteropm (dentition, as a rule, unusually heavy) 
there is a pronounced tendency to an enlargement of the cingulum, 
particularly in the upper canines, and m certain species {Ft. 
samoeasis and pselaphoa groups) the prominent edge of the 
cin-ulum tends to break up into a number of small, rounded, 
more or less incompletely separated tubercles. The extreme of 
this tendency is seen in the related, very heavy-toothed FteraJopex, 
in which the edge of the broad cingulum of the upper canines is 
distinctly cuspidate. 

Secondary cusps.— ^aj be developed from the inner edge only 
(in both upper and lower canines), or from the outer edge only (in 
the upper canines), or from both edges (very rarely, and only 
in the lower canines): — _ 

From inner edge only :— In Cynopteitis (fig. 48, p. o88) and the 
closely allied Ptenochirus (fig. 51, p. 643) a distinct secondary cusp 
is developed near the middle of the inner edge of the upper and 
lower canines, produced by a prolongation of the cingulum. 

From outer edge only:— A small, well-marked secondary cusp 
from the outer edge of the upper canine, above the middle of the 
tooth, in Pteropus tubercidatus : a similar, but much larger cusp 


halfway up the outer edge of the upper canine, in Fteralopex 
(fig. 21, p. 483); a similar, strong cusp in Harjiyionycteris (fig. 78, 
p. 800) ; often a more or less well-defined outer cusp in the upper 
canine of jYyctimene (fig. 61, p. 684). 

From hoth edges : — A strong secondarj' (cingulum) cusp at lower 
half of outer edge and a smaller one at middle of inner edge, in the 
lower canine of Harpyionycteris (fig. 78, p. 800). In this genus, 
it will be noticed, the upper canines are bicuspid, the lower 

Grooves. — The anterior (or antero-medial) surface of the upper 
canine is not infrequently marked with a deep and well-defined 
longitudinnl groove, reaching nearly from the base to the tip of the 
crown (and corresponding to the line followed by the tip of tho 
lower canine, along the crown of the upper canine, when the lower 
jaw is moved up and down) ; this groove is particularly con- 
spicuous in Boneia (as a rule obsolescent or shallow in the related 
liousettus), Pteropus (rarely obsolete ; in the related Pteralopex 
shallower or rather indistinct), Styloctenhim, in all genera of the 
Cynopterus section except Myonycteris, Cynopterus, Meyarops, and 
Nyctimene, and in all Macroylossince. More rarely, and only in 
some genera of Macroylossin(x, there are one or two additional 
longitudinal grooves on the outer surface of the upper canine 
(barely traceable in Macroglossus, shallow in Syconycteris, as a rule 
better developed in Melonycteris, Nesonycteris, and 2iotopteris). 

Direction. — The lower canines are slanted conspicuously outward 
or both outward and backward in Boneia, PleroU-s, Epomops, Eyp- 
siynathus (lower canines, like lower incisors, closing some distance 
in front of upper), Sphcerias, and all Maeroylossince. In Dolsonia 
the upper canines are a little proclivous ; in the related Harpyio- 
nycteris both upper and lower canines (as well as the incisors) are 
unusually proclivous, the lower canines crossing the upper almost 
at right angles. 

Position of lower canines. — In genera with a single pair of lower 
incisors the lower canines have, as a rule, moved a little closer 
together, toward the extremity of the jaw {Siyhctenium, Pteno- 
cMrns, Meycprops, Balionycteris, Penthetor; not in the two Macro- 
glossine genera Nesonycteris and Notopteris). The extreme phase 
of this tendency is seen in three genera, Bohsonia, Harpyionycteris, 
and Nijctimene, in which the lower canines have moved forward to 
the very extremity of the jaw, so as to be quite or nearly in 
contact with each other (in Dohsonia and Harpyionycteris the 
single pair of lower incisors is rudimentary, wedged in between the 
canines, and at least in Dohsonia sometimes deciduous ; in Nycti- 
mene all lower incisors are lost). 

11. Premolars and molars. 

Formulct. — p^ and p^ are lost in all Chiroptera, m' in all Mega- 
chiroptera ; no Fruit-bat, therefore, has more than ^ postcaniue 

xxvm pimstk; ciiakactkus. 

tfotli on ciUior side. The following seven formulae occur in tlic 
Buborder : — 

ji' p' p* m' m'' _ ''^ ii 

It n~T) nrm~ni ~ G' "ormal formula, found in all gcneni not 

enumerated helow. 
p' p' p^ m' m- _ 5 . 

11 i)~rrnr~m^^ ~ ^i' '" '^fll'-f^cteidani (related to I'icropus) ; Bal'io- 

njfckrts (Cy nop terns section); and one species of 
Macrojlossincn, Konj/cteris rosenberr/i (tlic other 
species of Eonycteris have the normal formula). 

— p' p' m' m^ _ 4 . ^ ■ ' 

^- _^ m Dohsoma, an aVjcrrant trenus of the 

P. P3 Pi ™i °i. ™.-, '^ 

llouscllus section. 

p'p3p4mi_ _4 . ,„ 

-, in y lerotes (hnomop/iorus section). 

Pi p.. Pi f"i "'j I",-, ^ V i i / 

])' p" p' m' — _ 4 ,, , „ , . , ^, 

,, the usual lormula la the CynopUrus section, 

I'i Pi Pi "^j "'■ — " J J. 1 

found in eight genera, viz. Cijno [items, PteiiocJiirus, 

Mcrj(erops, ChirorMX, Thoo/iterus, Penlhelor, Splurrias, 

and JVyctimene; further, in one sjjecies of Macro- 

(flossime, iSj/ronT/cteris nains (the other s[iecio3 of 

Syconijcleris have the normal formula). 

— 1)^ p* m' m* 4 . 

= in Kotorderls (subfamily Macroqlossinoi) . 

— P3 Pi ™> "1. ^3 5' ^ ^ J J J 

— ~ — = -, the usual formula in the Eiiomoi/honis sec- 
Pi P3 P4 "i. ™.— 5 

tion, found in seven genera, viz. Epoinops, Jfiji'- 

stf/iialhii8, Epomopliorus, Microplero/ms, Nanoni/c- 
leris, Scotonycteris, and Casinycteris ; and in one 
genus of the Cynopterus section, JJyacoplerus. 

Formuhj: in iJie four 2->rimary sections of MeyacJiiroplcra. — All 
genera of the Itovseltus section Lave the normal number of pre- 
molars and molars (''), except Styloctcniwa, which has lost the 
small nij, and JJohsonia, which has lost the small p'. All genera of 
the Epomophoriis section have lost p', m'', and m^ (^ cheek-teeth), 
except Fhrotes, which has lost only m% and retained p' and m,, in 
a rudimentary state. All genera of the Cynopterus section have 
lost m* and m^ (* check-teeth), except Myonycteris, which has the 
full numher of check-teeth (but m^ and m^ are quite small), 
Bulionyetcris, which has lost only m^ (but m' is minute), and 
JJyacopterus, which has lost not only m^ and m^, but also the small 
p'. All Jilacrofjlosshue have the normal cheek-tooth formula (j!)^ 
except Eonycteris rosenherrji, which has lost m,, fin the other species 
of Eonycteris m^ is usually very small, and adult individuals 
occasionally occur in whicli it is absent at least on one side), 


Si/con>/cte)-is iKiias, which lost in, and nr (hotli small in tlio 
other spt'cios of iho goims), aud JSotopta-ia, which lias lost p' and p 
(in the related MclonT/cteris and AcnoDi/citris p' is rudiiuoiitarv'). 

Modijications of molar sirwture. — Normally the structuro of the 
molaritbnn teeth iu Me?:achiroptera is this: a median lonjiitudinal 
groove flanked by a higher enter and lower inner ridge, eaeh ridge 
lising (or tending to rise) into a cnsp anteriorly. This typical 
structure is most clearly pronounced in p' and ni', p^, m,, and ni„. 
In p-' and p., the outer and inner ringo raises anteriorly into a 
higher, more narrowly pointed cnsp, and both cusps are either 
connected at base by a conspicuous commissuro or (very often) 
completely fused from base to tip, making these teeth iii i)rotile 
almost eaniniform. If present, ]>', p, and m, are nearly alwavs 
reduced iu size, and their surface structure more or less de- 

The principal modilkations of the typical molar structuro are 
due to : (1) the development of a well-marked posterior basal ledge, 
(2) the development of an antero-internal basal ledge or cusj)! 
(13) a more or less complete splitting of the outer or inner ridge, or 
both, into two or more cusps, or (4) the development of surface 
cusps. If only one of these modifications becomes operative, the 
general ai>pearauco of the tooth is comparatively little altered; if 
several (/'/crw/o/),;.!-) or all {JJarpi/ioni/dena) take effect in the same 
tooth, its appearance is naturally profoundly modilied. 

In rta-opus there is a pronounced tendencv to the developnu-nt 
of a posterior basal ledge in p', p', p.,, and p„'often also in m' and 
nip not infrequently in m.^, i.e. the posterior portion of the tooth is 
more or less distinctly marked off, by a notch in the outer ridge, 
from the rest of the tooth (fig. 9 B, p. 08; llg. 101], p. (Jil); these 
posterior ledges are unusually strong in the rurojnis samoensis and 
pseliqtJwn groups, and in one species of the former group (7'/. 
anedanus) the ledge is continued forward, as a well-defined shelf, 
along the inner side of p^ ni„ and ni.^ (fig. 10 C, p. (;9 ; fig. 14* 
p. 289). In some species of the Ptcro}}iis lomhoccush, samocnsis, 
and psilajthon groups there arc traces of antero-internal basal 
tubercles in \)\ p^ p„ and p, or in some of these teeth, i.e. the 
antero-internal cingulum is some\\hat more ditVcrentiated thuu 
usual and tending to rise as a small ledge or tubercle. 

Tliese modifications aro further developed and more definitely 
fixed in Jcerodon (closely related to rUropi.s) : the jiosterior 
ledges aro always strong, in p^, m,, and m.^ the ledge always extends 
along the inner base of the teeth as a broad, sh.irply-defined shelf, 
and a well-marked antero-internal tubercle is always developed in' 
])' and m', in some species also iu p' and p^, making these teeth 
distinctly tricuspid (fig. L'O, p. 413). 

The peculiar molar structure of PUraJojtcv (also related to 
rtcropux) represents only an extreme phase of the modifications 
developed in the sjiecies of the rtcropns jisclap/ion group. The 
teeth are shrM't and broad, the anteii(U- and post(-ri(>r ledges of }>", 


p\ and in' prominently developed, slielf-like, with raised margin, 
the posterior ledges of the lower teeth heavy, and the cutting-edge 
of the outer ridge of p^ and m^ distinctly (though never deeply) 
bifid (fig. 22, p. 438). 

Again, in Dohsonia (an offshoot from the Rousetto-Pteropine 
section) some of the modifications seen in Pteropiis are taken up 
and further developed. In a single species {Dohsonia minor) the 
molar structure is practically unmodified Rousettine, hut in the 
more highly diff'erentiated forms of the genus there is a well- 
marked posterior basal ledge in p^, p\ Pg, and p^, and a distinct 
antero-internal basal ledge or cusp in p' and p'', often also in p^, p^, 
m^, and m'; to these modifications are added, in the majority of 
species, a median surface ridge (or cusp) in m' and m^, sometimes 
also in p'' and m^, and a tendency to a splitting of both ridges of p*, 
m\ p^, and m^ (rarely p^ and ]}') into two or more cusps (fig. 25, 
p. 451). 

A distinct (but never very deep) splitting of the ridges of some 
of the cheek-teeth was noted above in Pteralopex and Dohsonia. 
The same tendency crops up in several other Fruit-bats : in 
Pteropus pselapJwn the inner ridge of m,, and in Pleropus leuco- 
pterus both ridges of m^ and m,^ tend to become bilobed ; in 
ffi/psignathus the outer ridge of p^ is more or less obscurely, that 
of m^ always distinctly trilobed or bilobed, that of m, bilobed 
(rarely, and only as an individual anomaly, an initial stage of a 
splitting of the outer ridge of m^, or both m^ and m', into two 
cusps is seen in the related EpomopJiorus ) ; in Nyctimene (Cyno- 
pterus section) the outer ridge of p^, p^, and m^ is sometimes more or 
less conspicuously bilobed ; and, finally, in Earpyionycteris (related 
to Dohsonia) the splitting of the ridges is more complete than in 
any other genus of the suborder. 

The occurrence of surface cusps (or ridges) in p'', m\ m^, and m^ 
of Dohsonia was mentioned above. Similar cusps are developed in 
the related Harpyionycteris, and in p, and m^ of four genera of the 
Cynopterus section, viz. Cynopterus (character not quite fixed), 
Ptenochirus, Dyacoptemis, and Thoopterus. 

The " multicuspidate " molar structure of Earpyionycteris is 
unquestionably the most peculiar in the suborder. In reality, 
however, it is eff'ected simply by a combination of all the four types 
of modification discussed above, viz. by the development of postero- 
external and antero-internal cusps or ledges and of surface cusps, 
and by a splitting of the ridges (see analytical description of the 
teeth of this genus, pp. 801-803, and fig. 79, p. 802). 

Deciduousness of pV — Even when reduced to a quite rudimentary 
(and therefore presumably almost functionless) condition p' is in 
most genera permanent. It is known to be deciduous in a few 
species of Ronsettus (seminudus, hrachyotis), in many species of 
Pteropus, and in Acerodon and Styloctenium (perhaps occasionally in 
Chironax). [It is permanently lost in Dohsonia, all Epomophorine 


genera except Plerotes, in Dyacopterus (probably ; only one specimen 
is known), and in Notopteris. The latter genus is the only Fruit-bat 
that has lost both p' and p^.] 

Degeneration of chceh-teeth. — Probably owing- to adaptation to a 
kind of food that requires little mastication, the cheek-teeth are 
imusually narrow (sublii)ear)iii >S'^6'no>M/c<er?s (subgenus oiRovsettus), 
a few species of Pigro/Jiw (particularly Pt. scapulatiis and ivoodfordi), 
Plerotes { Epomophorus section), Sphcprias (Cjfnoptervs section), and 
nearly all Macroglossina; (less so in Eoni/cteris and some species of 
Syconycieris than in the other genera). The narrowing of the 
teeth is not infrequently {Plerotes, many Maeroglossina;) accom- 
panied by a degeneration of their surface structure (flattening of 

12. Anomalies in the dental formula. 

Notes have been taken, during the preparation of this Catalogue, 
of any anomaly in the dental formula of the specimens examined. 
Altogether forty-four cases have come under observation, i. e. 
22 per cent, of the skulls examined exhibit individual aberrations 
from the normal number of teeth. There can be no doubt, how- 
ever, that for the large majority of Megachiroptera this estimate is 
too liberal ; the fact is that of the forty-four cases no less than 
twenty-three fall on forms distinguished by a more or less high 
degree of degeneration of the cheek-teeth, a condition which appears 
to be particularly favourable for the development of individual 
dental anomalies (see the paragraph "■ Odontonycteris," p. 754; 
the " genus " Odontonycteris was based on a Macroglossvs with a 
supernumerary molar on each side above) ; it would probably be 
approximately correct to saj^ that in any large series of skulls of 
Macroglossus, Eonycteris spelcea, and Ptero]>7is scapvlottis at least 
twelve per cent, will present anomalies in tlie dental formula, 
whereas in all other Fruit-bats taken together the percentage has 
been found to be only 0*9. 

The anomalies observed may conveniently be classed under the 
following four headings : — (1) Absence of teeth which are normally 
present in the species (it is hardly necessary to say that senile 
conditions have been left entirely out of consideration, and that a 
tooth has been considered " absent " only if there is no trace what- 
ever that it has ever been present in the individual), this anomaly 
in some cases " foreshadowing "' the permanent disappearance of the 
same tooth in related forms : (2) Appearance of teeth which are 
normally absent in the species, but present in related forms of 
Itregachiroptera : (3) Appearance of teeth which are normally 
absent in all Megachiroptera, but (since present in some Micro- 
chiroptera) have probably been present in some ancestral form : 
(4) Accidental outgrowths. Whether a case ought to be classed 
under the fourth or third category is, as noted below, sometimes 


Alisence of teeth that are normalli/ present in the species. — The moat frequent 
anomaly is the absence of the last lowei* molar (m, or ra^) or the last upper 
molar (ra'^}; more rarely p', pi, or i^ are undeveloped: — 

inj. — Absent on one side of the jaw, in four cases, viz. one Ptcropus vampi/r/f.i 
vampi/rus (British Museum, no. 9.1.,5.8(.)7), two Eonycteris spelaa 
(B.M.,, one Macroc/lossus minimus minimics (10.4.G.1.'»). 

Absent on both sides, in five cases, viz. one Pteropus qiqantcus (, 
four Pteropus soapwlatm (.')7. 10.24.1,, 8.8;8.'4, 

m,, is always greatly reduced in size in Pteropus and Eonycteris spelma ; 
in a gouus closely related to Pteropus, viz. Stylocteniam, the tooth 
has perniauently disappeared, and the same seems to be the case 
in one species of Eonycteris {E. rosenhcrgi). In Pteropus scapulatus 
and Eonycteris the cheek-teeth are tending to degenerate in size 
(breadtli), and in Macroglossus the degeneration is even more pro- 
m'. — Absent on one side, in one Pteropus scapulatus ( 

Absent on both sides, in one Macroglossus minimus minimus ( 

As noted above (under m^ both are forms with degenerated dentition, 
ni.,. — Absent on one side, in two cases, viz. one Cynopterus hrachyotis angulatus 
(, and one Cynopterus hrachyotis minutus (U.S. National 
Museum, 141243). In Cynop)terus and related genera the number of 

cheek-teeth is reduced to r, tliat is, nij and m^ are lost ; ui^ is therefore 

flie last lower molar, and it is always considerably reduced in size. 

p\ — Absent on both sides, in one Eonycteris spelaa ( In Eonycteris, 
as in many other Mogaehiroptera, p'^ is rudimentary ; in certain genera 
it is deciduous or lost. 

p,.— Absent on one side, in one Acerodon celehensis ( p^ is always 
small, but in no Fruit-bat deciduous, and only in one genus {Noiopteris, 
entirely unrelated to Acerodon) lost. 

ij. — Absent on one side, in three cases, viz. one Cynopterus hrachyotis angulatus 
(, one Cynopterus h. hrachyotis (, and one Cyno- 
pterus h.javanicus ( This anomaly, it will be noticed, has 
only been observed in Cynopterus, and ia several genera related to 
Cynopterus i^ is permanently lost. 

Occurrence of teeth that are normally absent in the species, hut present in re- 
lated Jorms : — 

lUj. — Present on one side, in one Cynopterus hrachyotis javanicus ( 
The tooth is normally lost in Cynopterus, but present in all Fruit-bats 
with unmodified dental formula. 

Occurrence of teeth that are normally lost in all living Mcgachiroptera, but no 

doubt have been present in some extinct form : — 

jn3_ — Present on one side, in five cases, viz. one Eidolon helvum (B. M., un- 
registered, specimen c', p. 15), one Rousettus lea/;hi (., one 
Bousettus seminudus (unregistered, specimen a, p. 39), and two Macro- 
glossus lagochilus lagochilus (, 
Present on both sides, in four cases, viz. one Eousettus agyptiacus (, 
one Pteropus gigantcus (, one Macroglossus minimus minimus 
(, and one Macroglossus lagochilus lagochilus (U.S. National 
Museum, 125316). 

\. n]3 is normally lost in all Megaohiroptera, but present in some Micro- 
cliir<)ptera. Of the nine cases recorded above of the occurrence 
of what seems to be an " m' " it is perhaps safest to eliminate the 
four observed in Macroglossus, owing to the nigh degree of degeneration 
of the dentition of that genus and the unusually frequent occurrence 
of supernumerary molars (compare the Marsupial genus Myrmecobius'.). 
There remain five cases in Eidolon, L'ousiitus, and Pteropus, and in ' 
view of the fact that all three genera occupy a low position in the 
suborder it is at least not unlikely that the anomaly is a reversion to a 
more primitive condition. 


Supermimerary teeth tliat arc certainly, or at least probably, merely accidental 
outgruwthi : — 

"13." — In one Pteropus vampynis skull ( a tooth is present, on 

both sides, behind i^ and leaning against the inner side of the canine. 

The possibility of this tooth being really an i^ (lost in all Mega- 

chiroptera, but preserved in some Microchiroptera) cannot of course 

be denied, but from the position and general shape of the tooth it 

appears more likely that it is an accidental outgrowth. 

i' reduplicated on both sides. — In one Maoroglossus mininuis sohrinus ( 

i- reduplicated on one side. — In one Cynoptenishrachyotisangnlatus ( 

" P2," /. e. a tooth occupying the position of a ]i^. — Present on one side, in three 

eases, viz. one Pteropus scapidatus (, one Epomophorus gamhi- 

aniis (Berlin Museum, 10171), and one Eonycteris spelcsa (, 

Present on both sides, in one EpomopJiorus yamhianus ( 

It would be interesting if this supernumerary premolar really repre- 
sented P2, a tooth lost in all Chiroptera, and, as in the case of 
"ia" (abovf), the possibility cannot be altogether denied. But it 
should be remembered that Pteropus scajndatus and Eonycteris are 
forms with somewhat degenerated teeth, therefore liable to accidental 
anomalies in the dentition, and the teeth of Epomophorus gamhianus^, 
though not exactly .degenerated, are remarkably small for the size of 
the bat ; further, that the diastema between p, and p^ is in all three 
forms considerably wider than elsewhere in the lower jaw, so that, if 
there is any latent tendency in the jaw to a development of a super- 
numerary tooth, this wide diastema so to speak " invites" it to crop u]) 
there ; and finally, that in two of the species, viz. Pt. fcopiilatus and 
E. gainliiairiis, cases are known (see below) of the appearance of a 
supernumerary premolar between p^ and p^ (compare a Eousettus 
angolensis with an abnormal tooth squeezed in between m' and m^). 
Tliis, to say the least, is a warning against too rash homologizations. 
Supernumerary tooth between p3 and p^. — On one side (tooth small) in one 
Epamophorus gamhianus ( 

On both sides (tooth well-developed), in one Pteropus scapulatus ( 
Supernumerary tooth between m' and m'. — On one side (tooth extremely 

narrow, abnormal in shape), in one Rousettiis angolensis ( 
" inj." — On one side, in four cases, viz. one Pteropus gigantens (]06.d), one 
Mctcror/lossus laqochilus nouns (, two Macroglossiis I. lagochilus 
(10.3i 23 ; L'.S. National Museum, 125316). 

On both sides, in two cases, viz. one Macroglossus mininuiS minimus 
(., one Macroglossus lagochilus nanus ( 

In no other geuus are anomalies in the cheek-tooth series so frequent 
as in Macroglossus (see p. 754). 

13. Palate-ridges. 

Eight ridges forming regular curves from side to side, the inter- 
dental ridges almost equidistant, the postdental a little more 
narrowly spaced, and some of the latter slightly notched at the 
middle, as if tending to split into a right and left half, such is the 
simplest, and therefore presumably the most primitive, type of 
palate-ridges known in Megachiroptera (see fig. 28, p. 484, the 
Epomophorine geuus Pleroten). A similar, more or less slightly, 
but never profoundly, modified arrangement of the ridges is seen in 
the large majority of genera of the Rousettine, Cyuopterine. and 
Macroglossine sections ; the number of the ridges may be a little 
increased, and, if so, the increase generally takes place chiefly on 
tlic postdental portion of the palate, the spacing of the ridges may 


l)e a little less regular, and some of the posterior ridges mf\y be 
more distinctly divided at middle (for those details see the descrip- 
tions and figures under each genus). But the P^pomophorine is the 
only section of Fruit-bats in which more essential modifications of 
the system of ridges occur; in that section, and in that only, nearly 
every genus has its own peculiar form and arrangement of the 
palate-ridges, and in some cases {Epomops, Epomojilwrus) even the 
species may be identified from the characters of their soft palate 
(or, if this is removed, from the impressions left by some of the 
heavier ridges on the bony palate). Omitting all details, the modi- 
fications of the soft palate in the Epomophorine section of genera 
are briefly these : — 

The simple palate-ridges of Plerotes have already been referred to 
above. In two of the three known species of the related Epomops, 
viz. E. franqueti and huettil-oferi (figs. 31 A, B, p. 489), the general 
arrangement is still comparatively little modified, but the three 
anterior (interdental) ridges are become thick and prominent : in 
the third species, E. dohsoni (fig. .31 C, same page) a similar thicken- 
ing of the three anterior ridges has taken place and, in addition to 
this, also the fourth and fifth ridges are modified, being heavy and 
triangularly prominent. The jialate of the closely allied Ni/psi- 
f/nathus (fig. 34 C, p. 504) differs only in less important details 
from that of Epomops franqueii. In Epomopiliorus (fig. 37, p. 516) 
all ridges (six in number, apart from a few thin and inconspicuous 
ridges at the extreme hinder edge of the palate) are thick and 
prominent. Micropteropus and Nanonijcteris exhibit the most 
peculiar modifications of the ridges in the whole suborder. Micro- 
pteropus (fig. 39, p. 556) in so far resembles Epomophonts (to which it 
is most probably closely related) as all the ridges (five) are thick 
and prominent, but the first ridge is typically hastate in form, with 
the point directed backward, the second to fifth divided by a deep 
groove extending along the median line of the palate, very broad 
in front and gradually n.irrowing posteriorly ; a glance at the 
palate of this animal is sufiicient to distinguish it from any other 
Fruit-bat. The latter remark would apply also to Nanonyctcrix 
(fig. 41, p. 561); the interdental ridges of this genus are rather 
similar to those of Epomops (probably one of its closest relatives), 
but the postdenfal ridges are increased in number, narrowly and 
very regularly spaced, thickened and elevated at middle (forming a 
j)rominent keel along the median line of the postdental palate), 
depressed and very thin laterall}- ; this is almost exactly the reverse 
of the type of palate-ridges found in Micropteropus, in which the 
soft palate is marked not with a prominent keel but Avith a dee]) 
and broad groove along the middle. Scotonyeteris and Casinycteris 
(fig. 45, p. 571) in so far recall Nanonycteris as the interdental 
ridges are thickened and the postdental ridges somewhat (Scoto- 
nyeteris) or much (Casinycteris) increased in number, but the latter 
are simple, thin, and serrate. 


14. Tov'jue. 

Tlio principal morlifications of the surface structure of the tongue 
have been described and figured on pp. 723-728, fig. 65. 

15. Wing-structaie. 

Detailed descriptions of the characters of tlie wing, in each genus 
of Megachiroptera, are given in the systematic part of this Cata- 
logue. The present paragraph intends to give only a general survey 
of the variahilit}- of some of these characters. [In all Fruit-bats 
there are three phalanges in the second finger, the terminal phalanx 
nearly always clawed, two phalanges in the third, fourth, and 

ThirtJ, fourth, and fiftli metacarpals. — There is never any very 
great contrast in the lengths of the metacarpals of the three long 
fingers. As a rule, however, the third is distinctly the longest, in 
some genera the fifth, while in others again all three metacarpals 
are practically subequal. The variations in this respect, within 
the four primary groups of Fruit-bats, arc briefly these : — 

In RousettHS and its closest relatives (Eidolon, Boneia) the third 
metacarpal is as a rule slightly longer than the fourth, which is a 
little longer than or subequal to the fifth ; the " indices " of the 
metacarpals (/. e. their lengths for a supposed leugth of forearm of 
1000) are in typical llousettus, respectively, 612, 5^5, and 586, in 
Eidolon 600, 668, and 641, in Boneia 671, 659, and 648. In 
Dohsonia (indices 621, 568, 586) and the related Harpi/ioni/cteris 
(697, 661, 673) the third has remained the longest, but the fourth 
tends to be a little shorter than the fifth. Finally, in Pteropus 
and its relatives, Acerodon, Pteralope.r, and Styloctenium, the fifth 
is slightly the longest, the third a little longer than or equal to the 
fourth (ex., Pteropus Jij/pomelanus with the indices 689, 670, 718, 
Pteralopex with 690, 659, 708, St>/locteninm with 723, 723, 739). 

In the Epomophorus section the third is nearly always slightly 
the longest, the fourth and fifth subequal (or the fifth tending to 
be the longer of the two) ; Epomophorus with the indices 680, 647, 
and 641, and Epomops with 728, 699, and 715, may serve as 
examples. In Scofont/ctcvis (682, 678, 688) and the related Casi- 
ni/rtcris (692, 683, 692) all three metacarpals are subequal ; and 
iri Phrotes (679, 698, 689) the fourth tends to be slightly the 

Again in the Ctjnopterns section the third is nearly always the 
longest, the fourth and fifth more or less subequal, though very 
often with a distinct tendency of the fourth to be the shortest, or 
the third and fifth may be subequal, the fourth slightly the shortest ; 
indices in Cijnopierus 640, 599, 625, in Bali any cterin 719, 697, 714, 
in yi/etimene 70S, 646, 678. 

Two genera of Macror/lossino' (and those two which also in 
skull and dentition are the least specialized in the subfamily), 
viz. Eon>/ctrris and Mer/aJoglossus, are nearlv Rousettine in the 


proportionate length of the metacarpals : third longest, fourth 
intermediate, fifth shortest ; indices in Eom/cieris 675, 656, 608, in 
Megaloglossus 760, 716, 670. In all other Macroghsstnce these three 
metacarpals are either practically subequal in length (Alacror/lossvs 
and Sycom/cferis, both closely interrelated), or the fifth is distinctly 
the longest [Melonycteris, Nesonyeteris, Notopteris, all closely inter- 
related) ; examples, Macrocjlossus 726, 733, 740, Nesonyeteris 760, 
755, 798. 

Phalanges of iJiird, fourth, and fifth fingers. — The second (ter- 
minal) phalanx of the third finger is always much longer than the 
first phalanx, but nearly always (for exceptions see below) decidedly 
shorter than the metacarpal of the same finger ; the two phalanges 
of the fourth finger are subequal, but with a very distinct tendency 
of the terminal phalanx to be the longer ; the two phalanges of the 
tlfth are subequal, the terminal phalanx being sometimes rather 
longer, but more often a little shorter than the proximal. In some 
forms, however, a conspicuous lengthening has taken place of the 
terminal phalanx of the third finger, making it subequal to or 
longer than its metacarpal ; this is the casein Eidolon, Slenonycteris 
(subgenus of liousettus), Boneia, Pleropus, Acerodon, Dohsonia, and 
Plerotes, and the three closely interrelated Macroglossine genera, 
Melonycteris, Nesonyeteris, and Notopteris. 

Subjoined is given in tabular form the absolute minima and 
maxima of the indices of the metacarpals and phalanges of the three 
long fingers, and for comparison the actual indices of one of tho 
shortest-winged (Rousettus) and one of the longest-winged (^Neso- 
vycteris) Fruit-bats. 

3rd finger. 

4tli finger. 

5th flnger. 



2 ph. 


1 ph. 2 ph. 



2 ph. 















Total length of fingers. — The third finger is always by far the 
longest of all ; next in length comes the fourth, averaging about 
four-fifths of the third ; then the fifth, averaging about eleven- 
twelfths of the fourth, in rare cases (Epomops, and a few 
Cj^nopterine genera) very nearly equal to, but never longer than, 
the fourth ; the second finger is as a rule somewhat more than 
one-third of the third ; the first one-half of the second, or a little 
more. The length of the hand is of course determined by the 
length of its longest finger, and to give a rough idea of tho varia- 
tions in the length of the hand as compared with the forearm the 
genera of Fruit-bats are arranged below according to the " index " 


(relative length, for a forearm of 1000) of the whole of the third 
linger (metacarpals and phalanges combined). This index is 
between : — 

luOO-1000 : — In typical Eouscttus (i. e. li. amplexicaudatus and 
allied forms), the shortest-winged Fruit-bats. 

lUOO-1700: — In the following genera of the Cjjno-plerns section, 
viz. Cynopteras, Plenochirus, and FentJielor. 

1700-1800: — (In the following genera of the liousettus section:) 
Eidolon (as a rule), Sttnonycteris, Bone'ui, and 
Dobsonia ; {Epomophorus section :) Eponiophorus, 
Micropteropus, iScotonycieris, and Casinycteris ; 
(Cijnopterits section :) Myonycteris, Meycvrops, 
Dyacopterus, Chiroiuuv, Thooptcrus, and Sphcerias ; 
{Jlucroylossince :) Eonycteris. 

1800-1000 ■.—{liousettus section :) Lissonycteris, Pteralopex, and 
JIarpyionycieris ; {EpomopJiurus section :) Flerotes, 
Eponwps, llyps'ujnathus, and Nanonycteris ; (Cyno- 
pterus section:) BaUonycteris ; (Alacroglossince :) 
Megaloylossus and Macroylossus. 

1900-2000 :—(A'ot«e^<((S section:) Pteropns (as a xxAo), Acerodoa 
(as a rule), and iStyloctenium ; {Cynopterns sec- 
tion :) Nyctimene ; {Macroglossince :) Syconycteris. 

L'()00-2100 : — In the three Macroglossino genera, Melonycteris, 
Nesonycteris, and JVotopteris. 

The table below shows the absolute minima and maxima of the 
indices of all live lingers, and for comparison the actual indices 
of one of the shortest-winged and one of the longest-winged 

let linger 
c. u. 

2nd finger | 

c. u. ' 3rd finger, 
(if present). 

4th finger. 

5th finger. 

Absolute minima 







Archceopteropus, — Archceopteropus transiens, Meschinclli, from 
the Upper Oligoceno of Italy, a bat which, judging from the length 
of its forearm, was equal in size to an Eidolon or a female 
JJypsiynathus, is the only Fruit-bat known only from a fossil state 
(on " liousettus " yatUardi, from the Middle Miocene of France, 
see footnote, p. 22), and as a tolerably complete impression of the 
skeleton of its fore-limb has been preserved, it may be of some 
interest to compare its wing-structure with that of living Mega- 
chiroptera. It must be pointed out in advance, however, that 
Meschinclli lias made a mistake in the identitication of two of its 


fingers. If hi^i identiiicalions M^ere correct, the indices of the third, 
fourth, and fifth fingers would be, respectively, 1486, 1450, and 
1046, i.e. the third and fourth lingers would l>e very short and 
practically subequal, the fifth enormously lengthened, a condition 
unthinkable except on the supposition that the form of the wing of 
Archcpopteropus was fundamentally different from that of any other 
bat, living or extinct. It is obvious that what Meschinelli takes to 
be the fifth finger is the third, and what he considers tlie third is 
the fifth. With this correction the indices of the metacarpals and 
phalanges of the three long fingers (calculated from Meschinelli's 
measurements) are as indicated in the bottom line of the following 
table : — 

3id flnger. ' 

4th finger. 

5th finger. 



2 ph. 



2 ph. 



2 ph. ^ 

Pleropodidcc, minima... 

„ maxima .. 
















And the indices of the total lengths of all fingers are these : 

, , „ ! 2nd finger ! 
let linger ^ ^ g^j g^gg^. ' 4th finger. 
*-■• "• ' (if present). ' i 


5th finger. 

Pleropodidrr, minima ... 303 i 608 1541 

,; m-jxima ...' 474 786 2071 

[Arch(Eopferopus S43 1 843 1946 




A comparative analysis of the hand gives the following 
results : — 

The first finger diilerod in no respect from that of living 

The second finger was somewhat less reduced in length (843, as 
compared with 608-786); as its first phalanx (inde.x 126) was 
very nearly similar to the average of recent forms (in which the 
index varies from 89 to 149), and the metacarpal (541), though 
unusually long, is equalled by that of the longest-winged living 
Fruit-bats (total variation 406-543), the greater length of the 
whole finger was due chiefly to the rather less reduced second 
and third phalanges (together 176, as against 64-162 in recent 
Fruit-bats, but the latter maximum index includes the claw, 
whereas in the fossil form only the claw iihalanx is included). 

The third metacarpal was conspicuous]}' shorter than the fourth, 
this again somewhat shorter than the fifth. Although in some 
living Fruit-bats the fifth distinctly tends to be the longest, so 
great a disproportion in the lengths of the three long metacarpals 


is not found in any recent Fruit-bat (but tliere are more or less 
close parallels in ilicrochiroptera). A glance at the first table 
above shows that the third metacarpal was quite ordinary in 
length, and the discrepancy due to the greater length of the fourth 
and litth. 

The first phalanx of the third finger was normal ; the second, 
as usual, by tar the longest of all phalanges, and relatively even 
somewhat longer than usual in Fruit-bats. The two phalanges 
of the fourth finger were nearly normal, though the first rather 
shorter than usual. The first phalaux of the fifth finger w^as 
normal in length ; but there seems to have been two phalanges 
distally to this (hence Meschinelli"s statement that the "third" 
finger, which in reality is the fifth, had three phalanges), and, 
curiously enough, these two distal phalanges together are equal in 
length to the single distal phalanx of living Megachiroptera (one 
might almost be tempted tn think whether these " two " distal 
phalanges are not one broken into two pieces, but it must be 
admitted that this suggestion is not borne out by the published 

As a rule in Megachiroptera the fifth finger is distinctly shorter 
than the fourth, though in a few genera (e. g. Epomops) practically 
equal to the lourth ; in Archceapteropus it was a trifie longer than 
the fourth (as in a few Microchiropteia). 

The general conclusions are : — In so far as the second finger had 
three phalanges, and its terminal piialanx was undoubtedly clawed, 
the hand of Archceopteropus was a genuine Megachiropteran hand ; 
and in so far as the second finger was less reduced in length, the 
hand of the fossil form may be said to be a little more primitive 
than that of any living bat : and in the features in which it 
differed, more or less slightly, from that of living Megachiroptera, 
it rather approached the hand of some Microchiroptera (except, of 
course, if it really had a third complete phalanx in the fifth finger). 
[The teeth of Archteopteropus are very little known, but the molar 
structure is said to have been cuspidate as in normal Micro- 
chiroptera, a statement that cannot be controlled with certainty 
from the published plate ; the tail was long as normally in 
Microchiroptera and in one genus of Megachiroptera, Noioj^teris.'] 

Claw of second fiiKjer. — The claw of the second finger is lost in 
one (aberrant) genus of the lionseltus section, viz. Do'>sonia, and 
in three genera of Macroglossinte, Eonj/cteris, Xesoivjcleris, and 
Aotopteris. Even if tlie claw is absent, the ungual phalanx is 
always present (rudimentary in Notopteris). 

Membranes. — The lateral membranes arise as a rule from the 
flanks or, rather higher up, from the sides of the dorsum. In 
Fteropm, Acerodon, and Stijloctenlum the line of origin lies 
generally somewhat nearer toward the spine, and in a few species 
of I'teropus {),ielunopo(jon, papuanus, ueohihet-nicus) the membranes 
arise verv close together, almost from the sides of the spinal tract. 


In Ptcralo^ex (related to Pleropus) they arise almost from the 
spinal line, or at any rate so close together that the interspace is 
scarcely appreciable. Finally, in two mutually entirely unrelated 
genera, Dobsonia {llousettus section) and Notopteris (Macroglossince) 
the naked membranes are perfectlj' continuous across the back and 
connected with the integument of the dorsum only along the spinal 

Posteriorly the membranes may be inserted on any toe, from the 
first to the fifth. The variations, within the primary sections, are 
these : — 

On first toe. — Itouseftns section : Eidolon, lioKsettus (pt.), Boneia, 
and Dobsonia (pt.), that is, in all genera except Pteropus and 
its closest relatives, and Harpyionycleris ; but in llousettus 
and Dobsonia the insertion varies between the first and 
second toe, and in Boneia the insertion is sometimes rather 
between the first and second toe than on the first. 
Epomophoms section : only in two genera, Scotonyderis and 
Casinycteris (in all other genera on second toe). Cynopterus 
"■ section : in Cynopterus, Ptenochirus, Meycerops, Balionycteris, 

PentJiefoi; and Spihcerias (in the other genera on second toe). 
Macroylossince : in Eonycteris either on first or second toe, 
or between. 

On second toe. — Rousettus section : in Pteropus, Acerodon, Plera- 
lope.v (sometimes on first), Styloeteniam, and Harpy ionycter is 
(perhaps rather between first and second); on Rousettus ?i,x\A 
Dobsonia compare remarks under first toe. Epomophorus 
section : all genera, except Scotonycteris and tkisinycteris. 
Cynopterus section : Myonycteris, Dyacopterus, Chirona.v (?), 
Thoopierus, and Nyctimene ; in the latter the insertion varies 
between the second and third toe. 3IacroyJossince: Eonycteris 
(often on first), MegaloyJossus (sometimes on third), Noto- 
pteris (sometimes between first and second). 

On third toe. — Rousettus section : none. Epomophorus section : 
none. Cynopterus section : sometimes Nyctimene (see under 
second toe). JIacroylossime : MegaloyJossus (on second or 
third), Macroglossus (on third or fourth), Melonyctcris (third 
or fourth), and Nesonycteris (third or fourth). 

On fourth toe. — Only \n sovae Macroglossince : Macroglossus {Wnrdi 
or fourth), Syconycteris (fourth or fifth, rarely between 
'■ third and fourth), Melonycteris (third or fourth), and 

Nesonycteris (third or fourth). 

On fifth toe. — In the Macrogiossino genus Syconycteris (fourth or 


The membranes are marked with well-defined yellow spots in 
Balionycteris and Nyctimene (both of the Cynopterus section) ; 
more or less obscure spots are detectable in a few other genera, 
e.g. Eonycteris. The wings are said to be "bright orange" in life 
in Casinycteris {Epiomophorus section). 


16. Tail. 
Varies ;is follows : — 

(1) Tailequul in length to forearm: — the oligocene ArchcFopterOjxis 
(8 free caudal vertebnc) ; one genus of living Megachiroptera, of the 
subfamily Macroglosshue, viz. Notopteris (" 10 vertebne,"' statement 
taken from literature, not verified by the writer). 

(2) Tail considerably reduced, subequal to or somewhat shorter 
than tibia, but longer than hind foot with claws : — one genus of the 
Cynopterus section, Xycthnene (7 free caudal vertebrae). Though 
the tail of Nyciimene is distinctly longer than in llonsettus, the 
number of vertebrae is the same as in Itousettus amplexiauidatus. 

(3) Tail further reduced in length, from about one-half to the 
full length of the foot with claws : — (i^owsc^^HS section:) Eidolon 
(4 free vertebrae), liouseltus (5-7), Boneia, Bobsonia; — {Cynoptenis 
section :) Myonycteris, Cynopterus (4), Ptenochinis, Dyacopterux, 
Fenthetor ; — {Macroylossimv :) Eonycferis (7). 

(4) Tail rudimentary, as a rule reduced to a small knob ex- 
ternally, more easily traceable by touch than by eye : — {Epomophorns 
section :) PJerotes (tail absent?), Epomops (2 free vertebrae), Epo- 
mophorus (2-3), Mkropteropus (3), Kanonycteris, Scotonycteris, 
Casinycteris; — (Cynopterus section:) Thoopteriis; — (Macroglosshia:) 
Megaloghssus (5 vcrtebrao, terminal two or three rudimentary), 
Macroglossus (3 or 2), Syconycteris. 

(5) External tail absent : — (Bouseti us section :) Pteropus (no free 
vertebrae), Acerodon, PteraJopex, SiyJoctenitim, Ilarpyionycteris ; — 
{Epomoplioriis section:) Hypsigmtthus (no free vertebrae); {Cyno- 
pterus section :) Megcerops, Balionycteris (no vertebrae), Chironax 
(no vertebrae), Spluerkis ; — {Macroglossinai :) Melonycteris, Xeso- 

In the first, second, and third of the stages recorded above the 
basal portion of the tail is included in (connected by its dorsal 
integument with) the intcrfemoral, the tip freely projecting (whether 
or not this was the case also in Archceopteropits is unknown) ; in the 
fourth stage the tail rudiment is usually unconnected with the 

Two facts are evident from the above, first, that the tail is con- 
siderably reduced in length in all living Megachiroptera, with the 
single exception of JS'otopteris, second, that the degree to which it 
is reduced varies, as a rule even very conspicuously, within each of 
the primary sections of the suborder. The Bousettus section falls 
into three natural subsections, the Rousettus, Pteropus, and Dob- 
sonia subsections ; in the first (Eidolon, Bousettus, Boneia) the tail 
is on the third stage of reduction, in the second (Pteropus, Acerodon, 
Pteralopex, Styloctenium) entirely absent, while in the third it is 
either on the third stage (Dobsonia) or absent ( Harpy ionycteris). 
In the whole of the Eponiophorus section the tail is rudimentary 
(fourth stage), except in HypsignatJius (and Plerotes'!), which has 
no tail. Within the Cynopterxis section, it is on the second stage 
in Nyetimene only, on (he third in Myonycteris. Cynopterus, Pleno- 
chirus, Dyacopterus, and Penthetor, on the fourth in Thoopleri'.!. 


and absent in Mcyicrops, Balioni/<;tei-is, Chlroncu', and Sphcerias. 
Finally, in the Macrocjlossinii;, it is on the first stage in Notopteris 
onh', on the third in Eonifcteris, on the fourth in Alegaloglossus, 
Mucrojlossus, and SyfonycUris, and absent in Melonycteris and 
JS'esoni/cteris, these two genera being the closest relatives of the 
long-tailed JSotojiteris. 

17. C'alcar. 

The calcar is rudimentary in Fhrotes (Ej)omo2^horus section) 
and Si/coni/cterls (subfamily Mucroghssitux), absent in Sphcerias 
[Cynoptenis section), the moditication being in all three cases due 
to a reduction of the lateral interfemoral ; in the two former genera 
the interfemoral is reduced to a narrow (Flerotes) or even sublinear 
(Syconycteris) rim along the tibia, in iSpha'rian it terminates at 
about the middle of the til)ia. 

18. CoJour of fur. 

In the majority of Fruit-bats the prevailing tinges of the 
coloration of the fur are a dark brown, sometimes inclining toward 
dark hair-brown *, sometimes closely approaching to bistre, or (not 
infrcqueutl}') washed with dull olive, the underparts being usually 
distinctl}' paler than tlie back. To have a basis for comparison 
this colour type may be considered the " typical " Megachiroptcran 
coloration, and the principal modifications, exhibited by genera 
and species differing from this " typical '' colour, may then be 
referred to one or several of the following categories : — 

(1) The dark brown tinge may brighten to fawn-brown, 
cafe-au-lait, fawn-drab or related tinges; or to wood -brown, 
yellowish, buffy or even cream ; or by increasing admixture of 
(sprinkling with) greyish hairs to pale hair-brown, ashy drab, or 
even light silvery grey (for examples see the brief summary of 
the colour changes in the i)rimary sections of Megachiroptera, 
below) : 

(2) The dark brown tinge may darken into seal-brown or 
blackish, either on the dorsal surface only (many species of Pteropus, 
some Acerodon), or both on the dorsal and ventral surfaces (some 
s])ecies of FtcropHs), this latter modification leading in its extreme 
to complete melanism (Pteropus modigUaiiii., Pt. natalis, some races 
of Pt. vampyrus ; Pleralopex ; occasionally in Pteropus alecto and 
Pt. tytleri) ; or a darkening of the underparts may be combined with 
a lightening of the colour of tlie dorsal surface (some races of 
Pteropus hypomelanus ; Melonycteris): 

(3) The fur of the nape of the neck (the " mantle ") may become 
brighter-coloured, forming a more or less strikingly-coloured 
(chestnut, russet, tawny, buff, yellowish buff, cream-buff, whitish') 
"tippet" contrasting with the unmodified or nearly unmodified 
dark back (many Pteropus, Acerodon) ; or the development of a 

* Throughout the whole of this Catalogue the colours are named, as far 
as possible, in accordance with Ridgway's ' Nomenclature of Colore ' (1886). 


poculiurly colouied mantle maybe conibiiied witli any of the coluur 
changes indicated under (1), (2), and (4): 

(4) The head may be marked with sharply defined stripes or 
patches (Ftcrojjus caimtraiiis and personahis, )Stylocteninm, Scoio- 
nycteris, Casinycleris) : 

(5) A small tuft of hair at the anterior and posterior bases of 
the ear-conch may become pale-coloured, whitish or bufty (eeere- 
ingly a trivial modification, but in fact eminently characteristic of 
all genera of the K2}onwp]iorus section, except one) : 

(G) A dark spinal stripe may develop (only in Nyctimene, and 
not equally distinct in all species) : 

(7) Peculiar light or bright-coloured neck-tufts may develop, 
the brightening of the colour sometimes spreading across the fore- 
neck and chest and backwaid along the fianks (in males of many 
species, rarely in females; see the- paragraph "Secondary sexual 
characters," below). 

In each of the natural sections of Megachiroptera the more 
noteworthy modifications of the colour of the fur are, briefiy 
summarized, these: — 

liousettus section. — Junistftus, Bnnna. and liarjiyionycteris are 
essentially '-typical'' in colour. In Eidolon {helvum and sabmim) 
the colours are more or less conspicuously tinged with yellowish 
(and "more so in females than in males). In Dohsonia the dark 
brown (Kousettine) general colour is often more or less brightened 
by admixture of olive, raw-umber, or tawny-olive, these tinges 
often with an indefinable greenish hue. P^eVojJMS (together with 
the closely related Acerodon) shows greater colour variations than 
any other genus of Fruit-bats ; any of the modifications mentioned 
above under (1) to (4) occur, single or combined; a summary has 
been given in the systematic part of this Catalogue (pp. 74-75). 
The t\yo known species of Pteralopex are nearly {aneeps) or quite 
melanistic (airafa). The single si>ccies o{ Styloc'tniimn is unusually 
light-coloured (silvery greyish or silvery biiflfy, with sharjjly con- 
trasting dark brown bases to the liairs), and with peculiar head 
markings, closely similar to those of Fteropus personatus, to which 
the genus is undoubtedly related. 

Epomophorus section.— Small whitish hair-lufts at the base of 
the ears anteriorly and posteriorly are present in all genera of this 
section, except iScotonyctcrin, but in no other Fruit-bats. The 
general colour of the fur is in Hypsir/nathus dull, dark ])lumbeous 
or slate, in all other genera as a rule more or less tending toward 
tho paler tinges of brown, fawn-brown, cafc-au-lait, brownish 
russet, brownish Isabella, or allied tinges. "White markings on the 
head, curiously analogous to those of Pteropus personatus and 
t'Stylocienium, are present in Scotonycteris and Casinycterix (the 
appearance of essentially analogous head markings in so widely 
separated forms as Pteropus personatus and Styhctenium on the one 
hand, Scofoiiyctei-is and Casinycteria on the other is difficult indeed 
to account for, except on the supposition that these arc cases of 


reversion to a colour pattern of the head of some common 
ancestral form). 

Cynopterus section. — Brownish tinges with paler underparts are 
the rule, except in Nyclimene. Myonycteris is practically llousettine 
in general colour. In Gynopterus the colour of the back is often 
suffused with warmer (russet) tinges, sometimes Avith cinnamon, 
or a tinge between cinnamon and wood-brown, the foreneck, sides 
of chest, and flanks with chestnut, cinnamon-rufous, or deep hazel 
(but generally much more so in males than in females). Mego'vops 
is somewhat paler above, approaching cafe-au-lait, like many 
Eporaophori ; Sphan-ias nearly greyish hair-brown. Balionycter'is 
and C7itVo?irtct' are rather darker above than usual, with the head and 
nape of neck nearly blackish. A well-marked dark brown spinal 
stripe renders most species of Nyctimene easily distinguishable in 
colour from all other Fruit-bats ; the stripe is generally narrow 
(one-fifteenth to one-eighth of the breadth of the back), sometimes 
obsolescent, sometimes again (^V. ucllo) very broad (one-third of 
back) ; the general colour of the upperside is sometimes irregularly 
mottled all over with darker tips to the hairs, as a rule, however, 
paler than usual in Fruit-bats, fawn-brown, fawn-drab, ashy-drab, 
■wood-brown, buff}', or even cream (the palest tinges seen in females 
of some species). 

Macrorilossince. — Eonyctcris is typically Eousettine in colour ; 
Megaloglossus and Notopteris dark. In Macroglossus and >Syco- 
iiycteris the colour of the upperside is lighter, varying from warm 
russet Front's brown to almost pure wood-brown. Melonycteris is 
approximately cinnamon above, with small white " epaulettes," 
nearly seal-brown beneath (an unusual contrast) ; the related 
Nesonycteris differing only by the pale underparts and the absence 
of " epaulettes." 

19. Size. 

To show their differences in size the Avhole series of genera of 
Megachiroptera are arranged below according to the lengths of the 
forearm (that this list can be only approximately correct is a matter 
of course, some of the genera being as jet known only from one or 
a few specimens). The smallest Fruit-bats are but little larger 
than the very smallest Microchiroptera, the largest greatly surpass 
in size any form of Microchiroptera. The " expanse " of a Fruit- 
bat is roughly about six times the length of the forearm (rather 
more than less). The extent of individual variation is in any Eat 
usually at least about ten, often twelve per cent., very rarely as 
much as fifteen to seventeen ; i. e. if in a sutficiently representative 
series of individuals, all belonging to one species or subspecies, the 
forearm of the smallest adult individual measures 50 mm., that of 
the largest will be about 55-56 or a little more, or if the minimum 
is 150, the maximum will as a rule not be more than 165, rarely 
as much as 170. 


37_ 42 mm.: — BaUonycteris; ? Chironcnv, jVer/alor/losxvit ; Marro- 

qlossiis ; Syconycteris. 
43- 49 mm.: — Nanonycteris ; Scotonycteris ; Chirona.v; ?Sph(f- 
rias; Megdloglossus ; Macroglossus ; Syco- 
50- 59 mm. : — Plerotes ; Micropteropns : Nnnonycteris \ Scoto- 
nycteris ; ? Caninj/cieris ; Myonycteris ; Cyno- 
pterus; MeyoTops ; Penthetor; Sjihcerias ; Nyctl- 
mene ; MeJovycteris ; Nesonycteris ; Notopteris. 
GO- 69 mm.: — Eousettux-^ Epomophorus •, Casinycteris ; Myo- 
nycteris; Cynnpterus; Penthetor; Nyctimene\ 
Eonyctcris ; Melonycteris ; Notopteris. 
70- 79 mm. : — liotcscttus ; Dobsonia; EpomopJiorns:, C'ynojiteriis ; 
Dyacopterus ; llioopterus ; Nyctimene ; Eonyc- 
80- 89 mm. : — Eousettus; Pteropus; Dobsonia; TImyyionycteris; 
Epomops; Epiomopilwriis ; Cynoptervs; P/eno- 
cJiirus; Nyctimene; ? Eonycteris. 
90- 99 mill. : — Pouseftus ; Boneia ; Pteropus ; Sfyloetemmn ; 
Epomops ; Epomopliorus ; Cynopierus. 
100-109 mm.: — Pteropus; Dobsonia; Epomops. 
110-119 mm. : — Eidolon; Pteropus; Dobsonia; ffypsiynatJius. 

[Archcvopteropus would belong here.] 
120-129 mm. : — Eidolon; Pteropus; Dobsonia; Hypsiynathns. 
130-139 mm. :—£'?c?o?on; Pteropus; Acerodon; Pteralope.r; Dob- 
sonia ; Hypsignathus. 
140-149 mm. : — Pteropus; Acerodon; Pteralope.r; Dobsonia. 
150-159 mm. : — Pterojms ; Acerodon; Dobsonia. 
160-205 mm.: — Pterojnts; Acerodon. 
206-220 mm. -.—Pteropus. 

20. Secondary sexual characters. 

In all or nearly all genera (in so far as both sexes are known) 
the canines average at least a little heavier and the zygomatic 
breadth of the skull at least a little greater in males than in females. 
In several Pteropodino' (some species of Pteropus; Pteralope.r; 
Penthetor) and the majority of Macroylossince (Macroglossus, Syco- 
nycteris, Melonycteris, Nesonycteris, Notopteris) this seems to be 
tlie only tangible secondary sexual differentiation, and even this 
difFerenco is in some cases so small as to be only appreciable in 
largo series of individuals. As a rule, however, the differentiation 
of the soxos is somewhat more conspicuous, the secondary sexual 
characters being referable to the following four categories : — 

(1) Males of a few genera average noticeably larger than females 
{Epomops, llypsignathus, and Epomophorus ; Eonycteris). 

(2) The general colour of the fur is in a few species conspicuously 
lighter in females than in males (some species of Eidolon and 
Nyctimene) ; or males average poraeNvhat richer in general colour 
than fomnles (Dobsonia riridis). 



(li) A tiift of hiiir on either side of the nock is in a large niimher 
of genera distinctly differentiated in the males, being moi'e rigid, 
unctuous, and brighter (or deeper or more saturated) in colour than 
the surrounding fur (development of " neck-tufts ") ; or the -whole 
of the fur across the foreneck and anterior portion of the chest 
may be more rigid and more saturated in colour in males than in 
females (development of a " ruff"), or the hair of the same region 
of males may differ, if not in rigidity, at least in its richer colour 
from that of females ; or (as is the case in some species of Pterojnis) 
there may hardly be any sexual difference in the fur of tlie fore- 
neck, but that of the nape of the neck (the " mantle") may be 
distinctly more rigid in males, and in some of these species pale- 
coloured from tip to base, whereas in females of the same species it 
is pale-coloured only at the exposed tip, dark-coloured at the con- 
cealed base {Fteropvs mariannus and conspic Hiatus groups). In 
some genera of the Epomopliorus section the " neck-tufts " of tlio 
males have, so to speak, moved higher np on the sides of the neck, 
being situated one on each " shoulder," i. e. the region of the sides 
of the neck immediately in front of the origin of the antebrachial 
membrane (" shoulder brushes" or "epaulettes,'' hence the name 
" Epomo-phorus," epaulette-bearer), at the bottom of a deep, pouch- 
like depression in the skin (" shoulder pouches), each shoulder 
brush being usually erectile and retractile at the will of the animnl 
("when erected the tuft had a vibratory movement," collector's 
note on the label of a Micropteropus pnsilhis, British Museum). 

(4) Helatively rarely (Epnmops, Ht/psirpiaiJiiis, Epomopliorus) 
males differ from females by the possession of one or several 
pharyngeal air-sacs, a peculiarity very often, perhaps always, 
combined with an enlargement of the larynx of the males (the 
voice of Epomops and J/j/psignntJnis (both sexes? or males only?) 
is described as a loud croaking, that of the latter i-ecalling the 
croaking of the Ethiopian Giant Frog, liana occipitalis). In one 
of the same genera {Hi/psignatlms) the muzzle and cranial rostrum 
are much heavier in males, owing to the development in that sex of 
a subcutaneous air-sac on either side of the muzzle and large 
cutaneous folds on the extremity of the muzzle (the latter present, 
but much smaller, also in females). 

lleviewed in each of the four natural sections of Megachiroptera 
the secondary sexual characters are, briefly, these (diiferences in 
size of canines and in zygomatic breadth omitted): — 

Rousettus section. — EiJolon : males with richer-coloured neck- 
tufts (traceable, but smalhsr and less rich in colour, in females) ; 
females, at least in two species (7ie?rM)n and sa/>«?Hm), conspicuously V 
paler than males, liousettus : males as a rule with neck-tufts, 
the unctuous hair sometimes forming a " ruff " across the foreneck. 
Boneia: sexual characters unknown (females not on record). 
Pteropus: fur of mantle often (not always) more rigid and unctuous 
in males, softer and more spreading in females, in a few species 


darker at concealed base in females than in males ; neck-tufts (not 
conspic-uous, except on spreading the fur) in males of some species. 
jicerodoti : essentially as Fterojjvs. Pteralopex: no appreciable 
differentiatio]!. Styloctenium: sexual characters imperfect.l}' known. 
Dobsonia : as a rule scarcely any appreciable differentiation, in at 
least one species (D. viridis) males distinctly richer in general 
colour. Harpy ionycteris: known from one adult specimen (unsexed). 

Epomophorus section. — Generally characterized by the unusually 
high development of secondary sexual differences. Plerotes: 
differentiation unknown (one female). Epomops : males with large 
shoulder pouches and erectile shoulder brushes ; males with two 
pairs of pharyngeal sacs (p. 492) ; larynx enlarged in males ; males 
averaging considerably larger than females. Hypsir/nathits : no 
shoulder pouches or shoulder brushes ; males with pharyngeal sacs 
and enlarged larj'nx, as in Epomop>s, and with a pair of sub- 
cutaneous rostral air-sacs; cutaneous folds on extremity of muzzle 
much larger in males ; muzzle and cranial rostrum of males much 
enlarged and differing in shape from those of females ; malts 
averaging much larger. Ej^iomophorus : males with large shoulder 
pouches and erectile shoulder brushes, and with one small central 
pharyngeal sac ; males averaging larger (character much less 
pronounced in the smaller than in the larger species). Micro- 
pteropvs and Nanonycteris : deep shoulder pouches and erectile 
brushes in males (females often with small pouches, but no brushes); 
scarcely any appreciable sexual difference in size. Scoionycicris 
and Cashiyderis : sexual characters uncertain (adult males un- 

Cyiwpterus section. — Sexual differentiation, if developed, usually 
Rousettine in character (though often more pronounced). 2Iyo- 
'iiycLeris: males with neck-tufts and " ruff " across foreneck (adult 
females unknown). Cynoj^terus and Pienochirus: males with ncck- 
lufts (small tufts sometimes traceable in females) and often with 
" ruff." Mi'i/a'rops and Dyaropierus : males unknot n. BaVto- 
nycterix : hair of foreneck brighter in males. Chirona.v r.nd 
Tliooptirus: adult males unknown. Penlhetor : no sexual difiVi- 
eiitiation. Sphccrias: males not seen. Nyctimcne: in some species, 
males with fur of sides of neck, foreneck, chest, and flanks much 
brighter in colour ; in other species, scarcely any sexual differ- 
entiation in the colour of the underpaits, but females much paler 
above (cream-buff, cream-white) than males (fawn-brown, ashy- 
brown, ashy-drab). 

Macroglossi7)a'. — Secondarj- sexual differences in most genera 
undeveloped ; if present, essentially llousettine, thougli in the case 
of one genus combined with a conspicuous average difference in 
size. Eonyciervi : males with a well-defined "ruff" across the 
foreneck of generally deeper (more saturated) colour, and averag- 
ing noticeably larger. Megalcylossrts'. males with large (whitish) 
neck-tufts. Macroglossns, Syconycteri/i, Melonycteris, ycsonycferis, 
and Aofoptervi : no conspicuous sexual differentiation. 


III. Interrelations of the genera of Megachiroptera. 

It has been considered desirable to give here a summary of the 
mutual affinities and probable phylogen)' of the genera, " sections," 
and subfamilies of living Fruit-bats, omitting as far as possible all 
discussion of details. For these latter the reader is referred to 
the paragraph " Affinities " under each genus, in the systematic 
part of this Catalogue. 

Rousettus (in its typical form, as represented by the species 
of the subgenus l^ousdUn-) is one of the least specialized genera of 
living Megachiroptera. The rostrum is moderate in length and iu 
no respect peculiarly modified, the premaxillse in simple contact 
with each other in front (not ankylosed together, nor spaced), the 
lateral margins of the postdental palate forming straight lines 
converging backward, the postorbital processes are short, the post- 
orbital foramina present, and the brain-case perfectly unmodified 
in general shajje. On the other hand, that all the most primitive 
cranial characters found in any living Fruit-bat should be united 
in the skull of this or any other living genus, is hardly to be 
expected; it is not difficult, therefore, to point out a few single 
skull characters in which Eouseitus stands a little higher than one 
or another genus of Megachiroptera : as in the large majority of 
Fruit-bats the infraorbital canal is quite short (considerably longer 
in MeJonijcteris, Nesonycteris, and Noto^^tens), and the ascending 
branches of the premaxilla3 as narrow at their upper extremities 
as near the alveolar margin (much less narrowed above in the three 
genera just mentioned); the palate is perhaps rather too broad to 
be considered quite unmodified in form, the tympanic bones, 
though typically annular, are rather broader than in most other 
genera, and the facial portion of the skull is distinctly, though not 
very strongly, deflected against the basicranial axis (more so than 
in the majority of Epomophorine and Cynopterine genera ; but it 
should be noticed that the deflection of the facial axis often varies 
to some extent in undoubtedly closely interrelated genera, and 
sometimes even in species of one genus : see anted, p, xxiii). The 
dentalformula is unmodified Megachiropteran (^ — ^ incisors, j. cheek- 
teeth), the teeth in no way peculiar, except in so far as the inner 
cusp of p' and p^ is completely fused with the outer, and that of 
p^ uearly so, characters shared with a large number of other genera. 
The palate-ridges are arranged in regular and almost equidistant 
curves over the whole of the palate, some of the posterior ridges 
always distinctly interrupted iu the middle; the tongue papilUe 
essentially unmodified (there is a faint tendency to a lengthening 
of the conical papillae at the extremity of the tongue : compare 
Macroglossina). A short tail is present, the second finger clawed, 
the wing-membranes inserted on the preaxial side of the foot, 
the prevailing colours of the fur a dark tinge of hair-brown or 
brown, the secondary sexual differentiation inconspicuous (males as 


a rule witli small, often more or less concealed, neck- tufts), and the 
general size moderate. 

The genus has divided into three branches (subgenera), Ronsetlu.', 
Sfenoiujclcris, and Lissom/cteris. The subgenus Eovsdhis (ten 
species) ranges over the Ethiopian and Oriental regions (extending 
from the former northward to Egypt, Palestine, and Cyprus) and 
the whole of the Austro-Malayan subregion, while SUnonycteris 
and Lissonjfcferis are confined to tropical Africa. In Stenonycteris 
(two species) the cheek-toeth are become narrow, m^ is reduced in 
size, and the facial axis more strongly deflected than in Eousettus 
s. str. Lissotij/cieris is somewhat more aberrant : the brain-case 
is peculiarly flattened posteriorly and the facial axis even less 
deflected than in Eousettvs, hoth characters giving the skull, viewed 
in profile, a rather striking resemblance to that of Epomops ; the 
prema.xillfe arc ankylosed together anteriorly, the postdental palate 
rejatively longer, the cheek-teeth shorter and broader (subsquarish) 
with the ridges more cusp-like (shorter antoro-posteriorly and 
higher vertically), those of p, separated, m' and p, reduced in 

Eidolon (three species, Ethiopian and jralagasy regions) has 
originated from a HouseUits-hke bat. p' is not quite so much 
reduced as in Roitseftus, but in other respects Eidolon is a little 
more peculiar : the premaxill?c are spaced in front, the postdental 
palate more expanded laterally, the tympanic bone elongated to 
form a short bony auditory meatus (a character unique among 
Bats),_ m, longer (antero-posteriorly) than usual, the sexual differ- 
entiation rather more pronounced (males always with neck-tufts, 
females often conspicuously paler than males), and the general size 
of the animals larger. 

Boneia (one species, Celebes) is closely related to liouKettu.t, 
but more specialized. The lower canines are slanted strongly 
outward, this necessitating a greater breadth of the palate between 
the upper canines, this again making the rostrum conspicuously 
broader in front; perhaps as a further consequence the premaxiila; 
are spaced anteriorly ; but in all other respects the skull is 
psscntinlly Rousettine in sha]ie. i' has disappeared, i^ is somewhat 
increased in size, the molariform teeth flatter, the outer and inner 
cusps of Pj separated and those of p' and p^ less completely fused 
than in liovseftus. The external characters are practically un- 

Pteropus, the largest genus of Megachiroptera (eighty-five 
species, a hundred and three forms) and one of the most widely 
distributed, has originated from a bat essentially similar to 
Rousettus, but slightly more primitive than the living species of 
that genus, in so far as it must have had a narrower palate, the 
tympanic ring thinner, and the cusps of p^ and p^ distinctly 
separated. The dentition is in all typical sjiecies of the genus 
considerably heavier than in liovseifus, hence the skull much more 
heavily built, the crests stronger, the postorbital processes lono^er, 



tl>o coronoid process of the mandible broader and more steeply 
ascending, i^ is always distinctly smaller than i^ (and in some 
species, chieHy of the Fterojms lomhocensis group, nearly rudi- 
mentary), p^ usually deciduous, m' and ni^ somewhat reduced ; the 
occipital (sublambdoid) jiortion of tlie skull is more elongaie, vub- 
tubular. The tail has entirely disapjiearcd, the colour's of the fur 
are mucli more varied (that of the nnpe of the neck, the " mantle," 
generally conspicuously brighter than the back), and the size of the 
animnls often greatly increased, tlio sjiecies varying in size from 
that of a .Fieldfare (or a medium-sized Jlousettus) to that of a Haven. 
Pteropus is distributed over nearly the whole area inhabited bj^ 
Megachiroptera, with the important exception of the African 
continent, a fact all the more remarkable inasmuch as the genus is 
relatively richly represented in the wliole of the Malagas}' region 
and occurring also in tiio island of Pemba, south of Zanzibar. 

The only claim of Acerodon (six species, nine forms: western 
Austro-Mitlaya and Philippine Islands) to stand as a genus distinct 
from Pteropus is the rather liigher specialization of its dentition : 
p'' and m' (sometimes also p^ and p.,) have develo])ed a well-defined 
antero-internal tubercle, and the lower molariform teeth (p^, m,, 
and m^) a sharply defined inner basal ledge; more or less distinct 
approximations to similar modifications are, however, seen in some 
specialized forms of Pteropus (see p. 415). 

The principal characters of Pteralopex (two species : Solomon 
Island^) are the excessively heavy dentition, the shortened, snb- 
sipiarish form of the upjier molariform teeth, the prominent anterior 
and posterior basal ledges of the same teeth, the bicnspidato outer 
lidge of the lower molariform teeth, the thick upper canines with 
a heavy external secondary cusp and small inner basal cusps, and 
the very large outer and small inner lower incisor. As pointed 
out in detail elsewhere (p. 436) all these peculiarities represent, in 
fact, onl}^ the last phase of modifications exhibited, more or less 
in an initial stage, by certain species of the Pteropus ^^sflaphon 
group (2'>sel'jpJwn, pilosus, tuhercuJatu?, and leucopterus). That 
Pternlope.v lias developed from a bat closely related to the living 
species of that group, or is, in other words, the peculiarly modified 
Solomon Islands representative of that group, is scarcely open to 

From another branch of Pteropus has developed the genus 
Styloctenium (one species: Celebes): i, and m^ have disappeared, 
and the colour of the fur is pale with dark bases to the hairs and 
sharply defined head markings,- — all characters so distinctly fore- 
shadowed in, or closely apy)roached by, the living species of the 
Pteropus iemmincki and lomhocensis groups as to indicate, almost 
with certainty, tlie origin of Sti/locienium from a form closely 
allied to the living species of those groups of Pteropus (for a dis- 
cussion of details, see p. 444). 

At some, probably not very remote, period of the growth of the 
Eousetto-Pteropine main branch of Megachiroptera a bat must 


have oxisfod which, though essentially similar tc a Pta-opu, i„ tlie 
Renenil appearance of the skull (palate narrower than in liou.ettn, 
occiput rather more » subtubular " than in nonsettu. but less so 
than in Ptcropus, tympanic ring thin) and with the whole series of 
cheek-teeth unmodified Eousettine or Pteropine in number ("_ •'*) 
nnd structure, differed in the following important dental characters'- 

rtnnlT'l ''"'r' .r''' '^'1^"J' P'-°^'i^-°^i-^ the lower canines 
Mtnated close together at the extremity of the mandible, tl>e 
nner pair of incisors above and below fi> and i, ) had disappeared, 
the Mngl. pair of lower incisors (i^) was quite small, the sinc^e 
pair of upper incisors (i^) somewhat modified in shape (crown 
obliquely triangular or obliquely bilobed, owing to it beini actid 
^.pon not by ,,, which was wedged in between the closely app'ox ! 
mated lower canines and much too small to reach i=, but by the 
tip ot the lower canines), and, as consequences of these modi- 
fi.'ations o the front teeth, the premaxilhx. were con iderablv 
reduced ,n breadth (sublinear) ; externally this bat must have beef, 
^cry hke a liovsettvs: tail i>resent (at about the length of 
l.e foot), second finger clawed, wings from sides of back aiK 
n.scrted posteriorly on first or second toe, colour of fur some dark 
IS 1 iT^Vn" ^^"-^^«r"' ^'^-^ P'-obably as a small lionsett..,. 
If not smaller still. From this bat have developed in one direction 

mZ:^. ^^"^^'•°-^^^'«^-)' -^ --t^.or//.,,,,/j„,,.,,,.(^MS!n: 

^.iL^t^Tt r^^^'K^ '^'"'''' ^''"■^'"" ^«'-"^^^'> the following 
further modifications have taken place: the small p> (which i^ 
sometimes deciduous in Eouseltru., and usually so in Fteropm) ha, 
entirely disappeared (cheek-teeth I), the wings arise from the spinal 
lino of the back, which is therefore completely covered bv the 
naked membranes, and the claw of the second finger has been lost' 
T. the most primitive species ot the genus (D. vvuor, Xew Guinea) 
the molar structure is typically Eousettine, but in all other spec cs 
it^ IS more or less complicated, by the development of posterior 
basal ledges antero-internal basal ledges, median surfacl ridges 
or by a tendency of the outer and inner ridges to break up into 
two or more cusps (for details see pp. 450-452 and fi- 9.5) ^ 

tvne ffltnlT^^^r"^ ^™"' ^^' above-described su^p^sed proto- 
t pe Harpyionycteris (one species) has been modified as follows- 
tt.e upper and lower canines and upper ir.cisor are stron^lv 
proc ivous (upper and lower canines crossing each other at neaHy 
r ghfc angles), tlie premaxilla, solidly fused anteriorly with each 
other and laterally with the maxilla^ an outer secondary cusp ha 
developed m the upper and lower canines, and a small inner^.usn 
m the lower canines (which therefore are tricuspidate), the number 
of cheek-teeth is unchanged (^: p' present), but the tendency in 

n^Inf rr'\°^- ^^'^'^-^^^^ (•"«« ^'>*-^-''«'«^ above) toward a develop- 
me^nt of posterior and anterior basal ledges and of surface cusL 
and to a splitting of the outer and inne^ ridges into two or more 



cusps has been carried to an extreme, so as to render the molari- 
form teeth raulticuspidate (for a discussion of the homologies of the 
cusps see pp. 801-803 and fig. 79). Externally, the onlj- noteworthy 
modification is the loss of the tail (wing-membranes normal, claw 
of second finger present). 

The nine genera reviewed above constitute what may be termed 
the Ronsettine section of Megachiroptera, in contradistinction 
to the Epomophorine, Cynopterine, and Macroglossine sections. 





Fig. II. — Interrelations of the gcner.a of the Eousetti/s section. 

It has divided into three branches (.sub.sections); the Rousettine 
branch {Ronsettus, Eidolon, Boneia), characterized by the simple, 
unmodified cranial characters (rostrum never shortened, premaxillae 
not sublinear, occiput not elongated, &c.), by the full Mega- 
chiropteran dental formula (except for the loss of i' in Boneia, and 
the occasional deciduousness of p'), the simple form of the premolars 
and molars, and externally by the presence of a tail and by having 
nearly always the third metacarpal slightly but distinctly longer 
than the fourth and fifth ; the Pteropine branch {Pterojius, 
Acerodon, Pteralopex, Stylo cteniuni) : cranial characters Rousettine, 
except for the more " subtubular " occiput and relatively narrower 



{jalute, dental formula unmodified (except for the loss of i, and m^ 
in ISti/lucteuinni), molar structure sometimes simple, but more often 
sliowing some, though rarely {Pieralope.c) a high, degree of special- 
ization, tail absent, and fifth metacarpal uearl}' always slightly 
but distinctly longer than third and fourth ; and the Dobsonian 
branch {Dohsonia, Harpy ionycteris) : rostrum at least somewhat 
shortened, premaxillse reduced in breadth, lower canines situated 
close together at the extremily of the mandible, i' and i^ lost, 
molariform teeth with a pronounced tendency to a high degree of 
specialization, tail present or absent, of the three long metacarpals 
the third nearly always distinctly the longest, the fourth the 
shortest, the tifih intermediate. The Kousettine branch (eighteen 
species) has spread over the whole area inhabited by Megachiroptera, 
except Polynesia; the Pteropine branch (ninetj'-four species, a 
hundred and lifteen forms) covers the whole of the same area, 
including Polynesia, bat excluding the continent of Africa and the 
Eastern Medite:ranean countries ; while the Dobsonian branch 
(thirteen species, fourteen forms) is confined to Austro-ilalaya and 
the Philippines. 

The probable mutual affinities of the genera of the Rousettine 
section are expressed in the diagram (fig. II.) on the foregoing page. 

The Epomophorine genus Plerotes (one species : Ethiopian) 
probably originated from a primitive Uousettus-\\]\Q type. Except 
lor the less of m'- its dental formula is typically Megachiropteran 
(iu all other Epomophorine bats not only m", but also p' and m^ are 
lost, and, as shown in fig. 28, p. 4S4, even in Plerotes these two 
teeth are so small as to be nearly functionless), and its remarkably 
simple palate-ridges are easily derived from those of Rousettus or 
rather from a form in this respect somewhat more primitive than 
the living representatives of that genus (see fig. 29, p. 485). But in 
other respects Plerotes is highly specialized : the palate is unusually 
broad, the cheek-teeth greatly reduced in breadth (almost Macro- 
glossino), the molars and last premolar flattened, with scarcely a 
trace of the usual cusp-like elevations, the lower canines slanted 
outward, and the tail and calcar absent or rudimentary. 

Somewhat similar lines of development have been followed by 
the related genus Epomops (three species, four forms : Ethiopian). 
Like many, if not all, other Epomophori it subsists chiefly on soft 
juicy fruits, the contents of which it draws out rather by suction 
than by mastication, and its lips, pharynx, larynx, cranial rostrum, 
dentition, and palate-ridges have been modified accordingly. The 
lips are full, pendulous, and highly expansible; the pharynx long, 
wide, and greatly extensible, communicating with the oral cavity 
by a very restricted aperture, the larynx spacious, with ossified 
walls, and supported behind by the expanded hyoids, acting as an 
" exhauster" during the suction; the bony palate is broad (though 
not broadened to the same extent as in Plerotes), the interdental 
palate-ridges thick and prominent (fig. 31, p. 489), the canines and 


])' aud p, tliiii and .sharply pointed, adapted to piercing the rind of 
the fruit and keeping it in position, while it is squeezed between 
the jaws and pressed upward against the interdental paLite-ridges ; 
the other cheek-teeth are ndatively feeble, p\ m'% and m^ have 
disajipeared ; r is often deciduous. 

Hypsignatlius (one species: Ethiopian) is undoubtedly derived 
from a form very similar to Ejioinopx, but probably with the post- 
dental pahite narrower. The rostrum is greatly enlarged, particu- 
larly in depth (and much more so in males than in females), the 
lower canines slanted more outward, p, more reduced, the ridges of 
the molariform teeth higher and their cusps more narrowly pointed, 
the outer ridges of some of the lower teeth more or less distinctly 
bi- or tricuspidate, the iiremaxillae aukylosed together in front, the 
muzzle thick aud truncate (■' Hamuierheaded Bats"), the u])per lip 
with prominent integumentary folds, the palate-ridges (fig. 34, 
p. 504) with relatively slight moditicatiuns similar to those of 

The origin of JS'anonifctei-is, Scotoiii/cieris, and Casiiij/rteris (all 
monotvpic and Ethiopian) is perhajis not (luitc clear ; that they are 
rather closely allied to each other there cannot be much doubt, and 
the probability is that this small branch is an offshoot from a type 
related to, but less specialized than, Pleroies and Ejwnwps. In all 
three genera tlie rostrum is considerably shortened, in Scotoni/cferis 
and Cusinycteris almost to the same degree as in Cynoptems. and 
the dental formula is in all tl e same as in Epomops (clieek-teeth ^ : 
p', m-. and m, lost). They differ from each other chiefly in the form 
of the bony jialate. In Nanonycteris the postzygomatic palate 
is unusually short and broad (fig. 40, p. /iGO), in Scotonycteris 
of normal length, with the lateral margins converging backward 
in straight lines (fig. 42, p. 564), while Casinycteris exhibits the 
unique ])eculiarity of having no postdeutal palate, the mcsopterygoid 
fossa extending forward very nearly to the level of the last molar 
(fig. 43, p. 569). The palate-iidgcs, though specially modified in 
each genus (see p]). 561, 565, 571), are without difficulty derived 
from an Epomnps-MkQ pattern. 

Like Epomops, Epomophorus (eight species, nine forms : 
Ethiopian) feeds chiefly on soft fruits, but in having adapted them- 
selves to this diet the two genera have followed to a certain extent 
different lines of development. In Epomops the rostrum and palate 
are broadened, in Epomophoms long and narrow ; in Epomops the 
postdental palate is broad and flat, i. e. essentially Eousettine in 
shape, in Epomophorus it is more or less deeply depressed posteriorly, 
with prominent palation rim ; in typical Epomops the anterior 
interdental palate-ridges are thick aud prominent, the postdental 
ridges simple (figs. 31 A, B, p. 48!J), in Epomophorus the postdental 
are practically similar to the interdental ridges, all being thick and 
prominent (fig. 37, p. 516). The dental formula and the characters 
of the teeth are the same in both genera. 

i>:Tnr.RELATio>'s oi GI:^•EK.^. 


Micropteropus (one species : Ethiopian) has jn-obabl) origi- 
nated from a type related to, but riiore primitive than, tiie living 
species of Epomojjhoriis (wiliunit the lengthening and narrowing of 
the rostrum characteristic of these). As in Ejiomophorus the post- 
dental palate is distinctly depressed posteriorly, with raised palation 
rim ; the palate-ridges are thick and prominent, but separated by a 
deep and broad groove along the median line of the palate (lig. iJ'J, 
p. 55G); the dental formula as in the majority of Epomophori ([>', 
m% and m^ lost) ; but the rostrum is as short and heavy as in 
Ci/nopterus, the interdental palate broad posteriorly, and the post- 
dental palate abruptly narrowed behind the anterior root of the 
zygomatic arches (fig. 38, p. 555). 

The eight genera reviewed above (Plcrotcs, Epomops, Ihjpsi- 
(jnathiis, JVanoni/cieris, Scotonijcte'ris, Caslinjcteris, Epohiopliorus, and 
Micropteropus) constitute the Epomophorine section of Eruit-bats. 
The principal differential characters, as compared with the llouset- 
tine section, may be briefly summed up as follows : — Dentition on 
the whole weak, p\ m'-^, and m^ lost in all genera except Plerotes, 
which has retained p' and m^ in a rudimentary condition ; molar 
structure perfectly simple, except for the nearly total degeneration 
of all surface structure in Plerotes and the splitting of some of the 
ridges in Ihipsignutlins ; number of incisors unmodified in all genera 
(i — "'), except for the deciduousness of i" in Epomops\ facial axis 
only very little deflected against the basicranial axis (except in 
Plerotes) ; brain-case distinctly flattened posteriorly (the same 
]>eculiarity is, however, seen in one type of the Rouscttine section, 
Lissoni/cteris) ; form of postdental jjalate highly variable (in the 
whole series of Megachiro])tera this is the only section that shows 
any great variation in this portion of the skull) ; palate-ridges 
simple only in Plerotes, in all other genera more or less highly 
specialized (the only section of Megachiroptera showing any great 
modification of the surface structure of the soft palate, in fact the 
only one in which every genus may be identified with certainty only 
by an examination of the palate-ridges) ; tail eitlier reduced to an 
inconspicuous rudiment, more easily delectable by the touch than by 
the eye, and not connected with the interfemoral, or absent ; small 
tufts of white hair present at anterior and posterior bases of the ear- 
conch in all genera, except Scotoni/cteris ; secondary sexual charac- 
ters often unusually highly developed : males with shoulder pouches 
and erectile light-coloured" shoulder tufts (" epaulettes '' ; exception : 
]/;/psignathus), sometimes with pharyngeal sacs (Epomops, Ihipsi- 
i/iiathits, EpomnphorHs). and not infrequently averaging considerably 
larger than females (Epomops, IJifpsigna'tJms, several species of 
Epomophorvs). The range of the section is strictly confined to the 
Ethioj)ian region. Only Epomophorus is distributed over nearly the 
whole of this region ; "the other seven genera are inhabitants of 
the " West African Province " (approximately synonymous with 
the Great West African Eorest Tract), from the Guinea Coast east 



to, or iu the case of one or a tew species a little be3'oiid, Victoria 
]Syun/,a, south to Angola. Eeiiguela, and Damaralaud. 

The genera of this section fall into three natural groups (branches, 
subsections) : — (1) The Epomops branch, tlie genera Plerotes, 
Epomops, and Hiiptsignathus (five species, six forms) : rostrum long, 
palate broad, postdental palate simple, at least some of the postdental 
palate-ridges unmoditied (except in Epomops dobsoni) ; (2) the 
Nanonycteris branch, including JS'anom/cteris, ticofom/cteris, and 
C'f(sint/cleris (three species) : rostrum much shortened, postdental 
palate highly variable, though never as in the third branch, at least 
some of the postdental palate-ridges unmoaitied ; (3) the Epomo- 
phorus branch, the genera Epumophorus and Micropteropus (nine 
species, ten forms) : postdental i)alate depressed posteriorly, rostrum 
varying in length, all palate-ridges modified. 

The probable mutual affinities of the genera of the Eporaophoriue 
section are expressed in the subjoined diagram (fig. III.). 






Fig. III. — Interrelations of the genera of the Epomophoms section. 

Myonycteris (four species: West African Province) has in many 
respects remained on the P^ousettine level of development, while in 
others it exhibits modifications approaching to those of Cynoptertis. 
The general external appearance, the dental formula, and the palate- 


ridges are (luitc or nearly as in Huuseltus, but tlie rostrum is con- 
spicuously shortened, the facial axis less detiected, ni^ and m" (last 
lower and upper moLir) leduced almost to rudiments, the orbits 
larger, the nostrils more piomiuent, and the calcar weaker. One 
species, inluibiting the island of San Thome, Gnlf of Guinea, is a 
little peculiar in some details of its dentition and ought to stand as 
a distinct subgenus (F/n/(jelis, p. 57'.*, fig. 47). 

Cynopterus (six species, si.xteen forms: Oriental region, east to 
Celebes and Timor) is further modilied : the rostrum is still more 
shortened, the facial axis more nearly horizontal, m^ and m" have 
disappeared, more or less distinct surface cusps are often developed 
in pj and m, (see p. 589 and fig. 49), the upper and lower canines 
have develo])cd a small supplementary cusp on the inner edge, the 
papillie on the inner side of the lips are increased both in number 
and siiic, the palate-ridges more crowded, thicker, and more sharply 
lirojecling, the nostrils more })roniinent, the tail a little more 
reduced, the colours of the fur more inclining toward the brighter 
tinges of brown, and the neck-tuttsof the males more conspicuou&ly 

Ptenochirus (one species : rhilii))iiiie Islands) is an only slightly 
modilied offshoot from the CipiojjiiVKs tj'pe (showing j)articularly 
close atfinities to the " J\'ia<lius " grou]) of that genus, a small group 
of species inhabiting the Malay I'ciiinsula, tSuniatra, !Xias, and 
Java, and characterized by the rather broader, more sub-squarish 
cheek-teeth and rather more shar|)ly developed surface cusps 
in p^ and m^) : i^ has disap[)eared (the tooth is in Cynoptenis 
smaller in bulk tlian i.,), i' is considerably sliortcned (in Cynopierus 
already slightly so), and there is a distinct vertical groove on the 
antero-medial surface of the ujjper canines (scarcely traceable in 

Megserops (one species : Malay reuinsula, ISumatra, Borneo) 
is derived from a bat very simitar to Cynopterus but without 
secondary cusps in the canines and without surface cusjjs in the 
lower cheek-teeth ; i^ is lost, i'' shortened, some of the lowei' cheek- 
teeth (p,, m,) are become broader, more subsijuarish in outline, the 
rostrum is much deeper in front, the nostrils rather more pioniinent 
(subtubular), and the external tail has disappeared. 

Also Dyacopterus (one species : Borneo) has originated from a 
primitive " Cy)iojitenis," without secondary cusps to the canines ; 
the cheek-feetli above and, particularly, below are shorter and 
broader, m' and m^ somewhat reduced, j), and ro, liave develojied a 
conspicuous rounded surface tubercle, p' is absent (or deciduous), 
the prcmaxilla! solidly ankylosed together in front, the postdcutal 
palate shoi'ter, the i)Ostorbital foramina (piercing bases of post- 
orbital processes) quite small, the membranes inserted on second 
(instead of first) toe, and the fur of the. body unusually short aud 
closely adpressed. 

In Myonycteris m, and m* are present, though reduced ; Balio- 
nycteris (one species : Borneo) has lost m^, but retained m'- in a 



nulimciitary state : in all other Cynoptcriiie genera both m., atul m" 
sire lodt. lu the mere dental fornnila, therefore, Bnlioni/cteris has 
remained on a slightly more primitive stage than all other 
Cyuopterine bats, except Mi/onycferis, but in other respects it is 
peculiarly specialized: i^ is lost, i" shortened, m' and m, somewhat 
reduced, some of the lower teeth (p^ and m^) subscpiarish, p^ has 
developed an antero-external basal lobe, the ixjstorbital foramina 
liave disajipeared, the tail is absent, the wings longer than usual 
and marked with sharj)ly defined yellowish spots, and the general 
size of the animal is unusually small. 

Chironax (one species) is the Javan representative of the Bornean 
Balioin/cteris. The two genera are strikingly alike in nearh^ all 
characters of the skull, in the general aspect of the dentition, 
as well as in external aj)pearance, "nhile at the same time some 
of the peculiar characters of Chironax are too important to allow 
it to be included in Bulionycteris : — -Balioni/cteris has retained a 
small m", in Chironax this tooth is lost ; in Balionycteris i^ is lost 
and i" shortened, in Chironax i, is present and i'' normal in length ; 
Balionycteris has developed an antero-external lobe in p", in 
Chironax the lobe is represented by a well-defined cusp ; in Balio- 
nycteris the premaxilla? are in simple contact anteriorly, in Chironax 
ankylosed together ; and the wing-membranes of Chironax are 
probably unspotted, i'rom this it is not ditficult to suggest the 
characters of the common ancestor of these two genera ; it must, 
have been a bafc very similar to Chironax, but with a small m'' and 
the premaxillffi in simple contact; in Java it developed into 
Chironax (m^ lost, premaxillae solidly fused), in Borneo into Balio- 
nycteris {nf retained, i, lost, i" shortened, the antero-external cusp 
of p^ enlarged into a lobe, wings spotted). 

The two closely interrelated monotypic genera TJtoopterus and 
Fenthetor may be presumed to have originated from a bat similar to 
Cynopterus, with the same number (^ — -) of incisors, the same 
number of cheek-teeth (t : m^ and m" lost), the same length of the 
tail, and the wing-membranes inserted on the first toe ; but the 
raolariform teeth, particularly p^ and m,, have been unusually 
broad, quadrate in outline, and their inner ridge very low, the upper 
canines grooved, both upi)er and lower canines without secondary 
cusps, there was no trace of surface cusps in any cheek-tooth, and 
the postorbital foramina had disappeared. This form developed 
in Western Austro-Malaya into Thoopterus : incisors unmodified, 
jij and ra^ with large surface cusfis, tail rudimentary, wings from 
second toe; and in Indo-Malaya into Penthetor : i, suppressed, 
i* shortened, no surface cusps in any cheek-tooth, tail of normal 
length, wings from first toe. 

Sphserias (one species : Burma) is probably derived from a 
primitive " Cynopterus," i. e. a bat with the dental formula and 
other characters essentially as in C ynoj>ter us, hnt without secondary 
cusps in the canines and without surface cusps in the lower cheek- 
teeth ; hut it is in certain respects more aberrant than any of the 


foregoing Cynopteriiie genera. The premaxilla? are proclivons, tlie 
iiicit<ors likewise and tiieir crowns peenliarly dilferontiated (triangu- 
larly pointed), the rostrum is ratlier less shortened than usual in 
Cynopteriiie bats and very thin and low anteriorly, the facial axis 
more distinctly deflected than usual in Cynopteri, the canines 
(particularly those in the lower jaw) slanted outward, and the 
cheek-teeth considerably reduced in breadth ; the odontoid papillae 
on the inner side of the lips are few and small (as in the lloiisettine 
section), the tail is suppressed, the interfemoral extends only to the 
middle of the tibia, and the calcar, therefore, has disappeared. 

The prototype of the highly peculiar genus Nyctimene (thirteen 
species: Austro-ilalaya and Australia) Avas ])robably similar to 
Ci/nopterus in all the essential characters of the skull, dentition, 
palate-ridges, and lip papillic, but without the secondarily acquired 
dental characters of the living species of Ci/nopterns (no inner cin- 
guluni cusp in the canines, no surface cusps in the cheek-teeth), and 
with the tail less reduced in length, even a little longer (though very 
likely not with a greater number of vertebra;) than in Ilouseltus. 
Developing into yj/ctlmene this ancestral form was roodified as 
follows : — The lower canines moved forward close together to the 
extremity of the mandible, and probably as a consequence of this 
all lower incisors disappeared (compare Dohsoniu and llariiyio- 
iiijcteris, both with the lower canines in a similar position, with i, 
lost and i, present only in a quite rudimentary state and occasionally 
deciduous) ; also i' disappeared (nomj)are the veiy conspicuous 
shortening of this tooth in several Cynopteriiie genera), so that i' 
became the only remaining pair of incisors above and below ; the 
nostrils were elongated into cylindrical tubes and the extremity of 
the rostrum moditied to act as a support of these tubes : the rostrum 
became unusually high (truncate) anteriorly, the premaxilloe deep, hut 
short (not reaching nasals), and firmly ankylosed together in front, 
and the extremity of the nasals was produced forward and downward 
in the middle ; ];>robably as a further consequence of the extra- 
ordinary- development of the anterior iiarcs, the posterior narial 
jiassage and mesopterygoid fossa became broader and deeper than 
usual (hence the slightly "pandurate" outline of the postdental 
palate). The wings are spotted with yellow (as in BaHoiii/cteris). 
The posterior circumvallate papilla is divided into two (fig. Go A, 
p. 725), and the cardiac portion of the stomach indistinctly differen- 
tiated (compare the alleged insectivorous habits of the genus). (Fur 
a more detailed discussion of the characters and afSuities of this 
genus see pp. 6yi-(i94.) 

The eleven foregoing genera constitute what may be called tho 
Cynopterins sectioa of Fruit-bats (thirtj'-one species, forty-one 
forms). The principal characters that link these genera together as 
a natural group are these : — ( 1) the rostrum is always conspicuously 
shortened, in all genera (except Mi/onj/cferis and Sphcerias) so much 
so that the distance from the orbit to the cxtrcmitv of the na?als is 


con:>ideiaLly less than the breadtli of the rostrum across the lachrymal 
foramina ; parallels or close approximations are found only in 
certain genera of the Epomophorine section (Nanonyctens branch 
and MkfojiUropus) and in some species of Bobsonia : (2) the facial 
axis of the skull is in all genera (excejtt Myomjcteris and Sjjhcerius) 
only very slightly deflected ; a similar inconspicuous deflection of 
the face is seen in tlie majority of E])omojihori, whereas in the 
llousettiue and Macroglossine sections the lace is always somewhat, 
and often much, moi'e deflected : (3) in Myumjcteris the lust molar 
lielow and above (m^ and m") are reduced almost to rudiments, in 
Balioni/cteris m^ is lost, in all other genera both m., and m", reducing 
the cheek-tooth formula from the typical ilegachiropteraii J. to -, a 
formula very rarely found outside this section of Fruit-bnts (it 
occurs in one species of Sifconycteris, and in Notoptcris, but in the 
latter case owing not to the loss of m^ and m- but of p' and p,) : 
(4) only Mijonyderishaa preserved the Rousettine aspect of the soft 
palate, in all other genera (in so far as the palate-ridges arekno«n) 
the surface structure of the soft palate is remarkably uniform, the 
ridges being prominent and crowded ; an exactly similar aspect of 
the palate is not found outside this section : (5) in all genera 
(except j\Ji/oni/cteris and Sphcericis) the odontoid papllte on the 
inner side of the lips are more numerous, larger, and more crowded 
than in any other section. 

There is a marked tendency in this section to — (1) a siipjiression 
of the inner pair of lower incisors (i^, which if present is always 
distinctly smaller than i ) combined with a reduction of the outer 

•2 Jl -1 -2 

pair of upper incisors (i") ; the incisor formula . — occurs iu 

a few other genera {Styloctenium, Ncsonycteris, Notoptens), but 
generally without a conspicuous reduction of i': (2) a develoj)ment 
of surface cusps in p^ and m^ [Oytwpterus, Ptenoilnrus, and Dyaco- 
pterus, but not in the related Meyarops ; in Thoopterus, but not in 
the related Fentlutor) ; a similar tendency is shown only bj' 
Bobsonia and Harpyionycteris : (3) to a disappearance of the post- 
orbital foramen (present and well developed in Myonycter'is, Cyno- 
pterus, Ptcnochirus, and Meyarops ; rudimentary in Byacopterus ; 
absent in Balionycteris, Chironax, 2'hoopteriis, PentJietor, Sj'hcerias, 
and usually in Nyctimene) ; the foramina are present and well- 
developed in all other Megachiroptera witliout exception: (4) to a 
shortening or complete suppression of the tail. 

lixcept for Myonytteris, which is Ethiopian (West African), the 
section is confined to the Oriental region and Austro-Malaya, one 
genus [Syctimene) extending from the latter subregion to Australia. 

The question whether this section, so far as our present know- 
ledge of the genera goes, is perfectly homogeneous, i. c. includes 
only genera which are more closely related to each other than to 
any genus outside the group, must, in the opinion of the writer, be 
answered in the affirmative, but it is quite possible that some 
systematists would be inclined to consider the status of Myonyctcris 



a little doTibtful. The fact is tliat, this genus has retained nian^- 
characters of lioiiseUns, while in practically all the features in 
■which it differs from liousetttis it more or less closely approaches 
to Cynopterus. Whether a genus exhibiting characters of this 
description ought, in a linear arrangement, to be classed at or near 
the end of the Rousettine section or as the "opening" genus of 
the Cynopterine section, must necessarily remain a matter of 

With regard to the principles for the arrangement of the genera, 
only a few remarks are required. Putting for a moment J/yo- 
nycteris and Nyctimene at one side, it would be easy to divide all 
the remaining nine genera of this section into two groups accord- 
ing to the number of incisors (|— | or ^— ^ ) ; but. as pointed out 
elsewhere (p. 649, footnote), an arrangement based primarily on 



section ] 

Fig. IV.— Interrelations of the genera of the Cynopterus section. 

this character would be thoroughly artificial ; it would separate 
Ptenochirns and Megrn'ops from their closest relative, Cynopterus; 
Bdlionycteris from its Javan representative, Chironcuv ; and Pen- 
thttor from its eastern representative, Thoopterns; there is no 
doubt that the loss of i^ is a character developed independently in 
difl'erent branches of the section. The diagram above (fig. IV.) is 
based, therefore, on the following considerations : — .1///oh//W<'/-/<? 


may at once be se])arate(l, owiiij; to its less slioitcnorl roslruin and 
the presence of in,, and m" ; next the two closely interrelated 
genera Baliouj/cteyia and Cliironax, the former of which has f)rc- 
served m" ; as a tliird branch Nyctimene, and as a fourth Sphcpriax, 
on account of their peculiar specializations in diH'orent directions ; 
the remaining six tjenera may bo divided into two groups, those 
with {C'l/nopterus, Ptenochirus, 3Ier/a;rops, Dyacoptenis) and those 
■without jiostorbital foramina (TJiooptcrus, PentJufor), Dyacoptenis 
being in this respect intermediate (foramina present but minute). 

All the genera reviewed above, the Housel tine, Eporaophorine, 
anil Cynopterine sections, form together a Subfamily ot Mega- 
chiroptera, the Pteropodinge. The characters of this subfamily, 
in contradistinction to tlie second primary subdivision of Mega- 
cliiroptera, the subfamily Macroglossinse, have been fully discussed 
in the systematic part of this Catalogue (i)p. 723-728, fig. 65), 
and it is sufficient here to emphasize the fact that the only real 
difference between the two subfamilies lies in the greater special- 
ization of the tongue and the tongue papillie in the Macroglosshuv 
(adaptation to a diet consisting, at least partly, of pollen). 

Eonyctsris (three species : Indo-China, Indo-^falaya, Celebes) is 
on the whole the least specialized genus of JJacroylossinre. In the 
general characters of the skull, dentition, and palate-ridges, as 
well as in external appearance (except for the loss of the claw of 
the index), it is remarkably like lioHsetiihi, but the cheek-teeth 
are narrower than in typical liousettus, m., and m" somewhat 
reduced (m, absent in one species), the tongue is, of course, 
typicall}' Macroglossine, there is scarcely any trace of odontoid 
papillae on the inner side of the lips, and the difference in length 
l)etween the third (longest) and fifth metacarpals is greater than in 

Megaloglossus (one species: Ethiopian) is undoubted]}' derived 
from an Eonycteris-W^Q type, but is considerably more advanced 
in specialization. The molars are become low and linear, but p^, 
])\ and p., are practically unreduced in height (fig. 71 B, p. 74P), 
the rostrum longer and slenderer, the premaxillai ])roclivous and 
ankylosed together in front, the extremity of the mandible elon- 
gated, the tongue somewhat lengthened, and the tail rudimentary 
(in some individuals hardly traceable externally). 

Macroglossus (two species, six forms : Indo-China, Indo- 
jVFalaya, Austro-iLalaya) is the eastern representative of the 
Ethiopian Meyalor/lossus. It has evidently originated from a typo 
essentially similar to lUer/aloijlosnHS, but with p\ m^, and m^ less 
reduced in size; and it has carried the specialization considerably 
fiirtlier than Mfr/aloylossus. All cheek-teeth (p*, p\ and p., not 
excepted) are become low and linear (fig. 71 C, p. 749), the pre- 
maxilla) are even more proclivous, the mandible even more length- 
ened anteriorly, the mandibular symphysis longer, the facial axis 
much more strongly deflected, the third metacarpal shortened and 


the fiitli lengthoiieJ (bo as to mako the thir.l, fourth, and liCth 
metacarpals ec^ual in length), and the caudal vertebra; further 
reduced in number. 

Syconycteris (three species, seven forms : Austro-Malaya and 
Australia) is ilosely related to Macrotjlossus, but all upj)er and the 
outer lower incisors are cons])icuously enlarged and proclivons, 
the posterior molar above and below reduced (absent in one species), 
the interfemoral unusually narrow, and the calcar rudimentary. 

In the foregoing four genera of Macror/lossiiia', which rna}' be 
called tlie Eonycterine section, as well as in all Ptero/)odin(v, the 
infraorbital canal is short (the outer wall a narrow bridge of bono 
and its anterior aperture, the infraorbital for;imen, situated verti- 
cally below or closely in front of the orbit), and the premaxillte are 
not or only very little broader above (at nasals) than below (near 
alveolar border). In the remaining three genera of Macrofflossinc^, 
the Notopterine section {Meloni/cferis, Ncsom/cten'^, and iVoto- 
pteris), the infraorbital canal is much loss reduced, the infraorljital 
foramen therefore situated a considerable distance in front of the 
orbit, and the premaxilla? are less narrowed in their upper halves, 
being about thrive or twice as broad above as below ; and the 
species of one of these genera (To^o^j^er/s) are the only living 
Fruit-bats in which the tail has remained long. Externally these 
two primary sections of Macroglossina' may easily be distinguished 
by an examination of the wings: in the Eonycterine section either 
the third metacarpal is distinctly longer than the fourth and fifth 
or these three metacarpals are subequal in length, and the terminal 
phalanx of the third finger is always conspicuously shorter than 
the third metacarpal ; in the Notopterine section the fifth meta- 
carpal is the longest, and the terminal phalanx of the third finger 
subequal to or even longer than the third metacarpal. 

In point of dentition Melonycteris (one species: Xew Guinea 
RTid Bismarck Archipelago) is the least modified genus of the 
Notopterine section. The dental formula is typically Megachi- 
ropteran (incisors r, — ;„ cheek-teeth "!), all the cheek-teeth are much 
reduced in size and sublincar in form. The external characters 
are not peculiar, except in so far as the tail has disappeared (and 
the ventral surface of the body is much darker in colour than the 

Nesouycteris (one species : Solomon Islands) is the slightly 
modified eastern representative of MeloiufcWris : i, is suppressed 
and the claw of the second finger is lost (the ventral surface of the 
body, as usual in Jlegachiroptera, paler than the dorsal). 

In having retained a long tail, much longer than in any other 
Fruit-bat, Notopteris (two sjiecies : Western Polynesia) is more 
primitive than Mrlonj/cterii and Nesoi^ijcliris, in other respects it 
has to a certain point followed similar lines of development, but is 
considerably more highly specialized, p' and p. arc lost (the latter 



tooth ia in the two related genera quite small, the former nidi- 
mentary and almost functionless), p^is enlarged and moved forward 
flosely behind the canines, i^ is lost (as in Nesoyiyctcris), and i' is 
deciduous ; the extremity (premaxillary portion) of the rostrum is 
elongated, the corresponding portion of the lower jaw peculiarly 
hroadened, and the premaxillse solidly united in front ; the claw of 
the second finger has disappeared (as in Nesonycteris) and even the 
claw phalanx is reduced in size, and the wings arise from the 
spinal line of the back, which is therefore covered by the naked 

The Macror/lossimv (seven genera, with twelve sjiecies or twenty 
forms) are essentially Austro- and Indo-Malayan in distribution ; 
one genus (Macror/lossvs) extends westward into the Indo-Chinese 
subregion, another {Notopterls) is peculiar to Western Polynesia, a 
third (Si^coin/cteris) extends southward to Australia ; and one genus 
{Megaloglossus) is confined to West Africa. The subjoined diagram 
(fig. V.) illustrates the probable interrelations of the genera. 


o iB 

Macroglossus Syconyctsris 

u §. Megaloglossus 


Fio. V. — Interrelations of the genera of tlie subfamily MacrogloasiiKP. 

As pointed out above, this subfamily falls into two natural sec- 
tions, the Eonyctcrine and Notopterine. Tlie less shortened infra- 
orbital canal and broad upper extremities of the premaxillas in all 
Notopterine genera, and the ]iresence of a long tail in one genus 
of the same section, all primitive characf.ers lost in all other Fruit- 
bats, are evidence that the Macrof/Iossime as a whole, in spite of 
the often very high specialization in other directions of the living 
genera, are of slightly lower origin than the Pteropodincp (see 
fig. VI., next page). 




Fig. VI. — General view of the interrelations of the s<iibfamilies, 
sections, and subsections of living Megachiroptera. 

IV. Geographical distribution of Megachiroptera. 

1. Distribution of genera. 

In the table below the ciphers in the left-hand column, after 
each genus, if single indicate the total number of species in the 
genus, if double the first is the number of species, the second the 
number of recognizable forms. In the other columns is given 
tlie number of species of each genus known to occur in each zoo- 
geographical region or subregion (as defined by Wallace), and if 
the number of " forms " is ditl'ereut from that of species it is added 
between parentheses. (If the ciphers in one horizontal row are 
added together, the total sum will not infrequently bo found to be 
greater than the number of species or forms given for the genus ; 
the reason is, of course, that in such cases one or several forms of 
the genus extend through more than one region or subregion.) 
In the column headed "Ethiopian region" is included the single 
species (lioiisettus ce(/i/ptiacus) which extends into the Eastern 
IVIediterranean subregion (Egypt, Syria, Palestine, Cyprus). For 
comments on the table, see the paragraph " llemarks on the 
geographical distribution," pp. Ixxv-xcii. 



Rouse fills section. 

Eidolon (3) 

Kousettiis (14) ] 7 

Boneia (1) 

Pteropus (85, 103) 
Aoerodon (tj, 9) ... 

Pteralopex (2) 

Stvlocteniiiin (1) ... 
Dobsonia(12, 13)... 
Harpyionjcteris (1) 

Epo'inaphorus section. 


Eponiops (3, 4) 

Hypsigimthiis (1) 

Epomopborus (8, 9) 

Micropteropus (1) .... 

Nanouycteris (1) 

Scotonycteris (1) 

Casinycteris (1) 

Cynopterns section. 

Myonyeteris (4) 

Cynopterns ((i, 16) .. 

PtenoeliiruB (1) 

Megserop.s (1) 

Dyacoptenis ( 1 ) 

Balionycteris (I) 

Ohironax (1) 

Thoopterus (1) 


Splijerias (1) 

Nyctiuiene (13) 


Eonycteris (3) 

Megaloglossus (1) 
Macroglossus (2, fi) 
Syconyctoris (3, 7) 
Melonycteris (1) ... 
Nesonycteris (1) ... 
Notopteris (2) 


■;; c 







M o 





a i, 


g -- 

s - 

























14 (24) 


36 (39) 


12 (13) 




J (4) 

























The number of genera, species, and forms represented in each 
region and subregiou is as follows : — 

E. Mediterranean subregion ... 

Etiiiopian region 

Malagasy rejiioii 

Iiiflian & Ceyloiiese siibregions , 

ludo-Cliinese subregi<iii 



Auslraljiui coiiUneiit 















2. Distrihulioii of specks and suhspecie^. 

The subjoined j^cographical list of the species and stibspecics is 
based almost eutirel}' on the material examined by the writer 
during the preparation of this Catalogue. An asterisk after a 
generic, specific, or subspecific name indicates that the genus, 
species, or subspecies is not known to occur outside the area dealt 

Eastern Mediterranean Subregion. 

Egypt, Syria, Palestine, Cyprus : — IiousctOis at/t/ptiu£HS (also in Ethiopian 

Ethiopian Region, 

West African Province (Great West ACrican Forest Tract, south to Daniara- 

lanil, east to Victoria Nyanxa ; one of the forms marked aa autochthonous 

extends a little beyond the eastern limits of the proyiiioe) :^ 
Eidolon /;cVw(;/J (Ethiopian region generally). 
lioitsettiis acji/pliactts (N.E. to Egypt. Syria, &c.) ; lanosus* (Ruwenzori) 

aiif/oleiisis * ; soii/hi* (Guinea coast). 
I'lero/es* aiich/c/ce (Benguela). 
Eponwps* jrauqufti sirfijifaiis (Gold Coast, Lagos, Ts'igeAa.); fro liquet i 

fraiiqucti (C)ld Calabar to Bukoba); hueitVcoferl {Sierra, Leone, Liberia) ; 

doluvni (Beugiieln, Katanga). 
Nifpsiffiift/fius * 7>ion.-itro.''i(s (Gambia to Monbuttu). 
J'jiij/iiiijjhonis wahfhet-gi halih'miuu* (Canieroojis to G. & B. East Africa) ; 

(;n/?(A;Vrw?^< (Senegal to Sennaar and Abyssinia); angulciufis* (Benguela, 

Damaraland); puuaarffuefi * (Upper Shari). 
Mici-op/erupiis * ptiHllus (Gambia to Victoria Nyanza). 
y<iiioiii/cferis * veld/iampi (Guinea coast). 
JSii:f()4ii/cferis * :eiikcri (Cameroo7is, Fernando Po). 
CiiMini/r/er/s * aryynnis (Can>eroons). 
Mi/o'ii/cfcris* wrimt/h/oni (Welle district); lepfodon (Sierra I^eone, 

Lil)eria) ; turquata (Lower Congo, Angola) ; braelii/cejtkala (Sau 

Mi'tjaloglossus * woermanni (Congo to Liberia). 

Eastern side of Continent : — 

Eidolon hclviiiu (Ethiopian region generally). 

liouscitus leachi* (B. East Africa to Cape Colony) ; rP(7.w7V?V?f«(Erylhrra, 
south-west to Gaboon and Loanda, norlli t(> Egypt, ■Syria, &c.) ; Kcrnpi * 
(Shoa, B. East Africa). 


Epomaphorws waklbergi wahlbergi * (B. East Africa to Cape Colony) ; 
lahiatus * (Sennaar, Abyssinia, Shoa) ; minor * (Shoa to G. East 
Africa) ; anurm * (Abyssinia aud Erytlirea to Tanganyika) ; ci-ypturus * 
(Nyasa to Transvaal); yambianus (Sennaar and Abyssinia, west to 
Guinea Coast). 

South Arabia : — 

Eidolon sahcsiMii *. 

Uoiisetfus arahicus (also Karachi). 

Peuiba Island : — Ptcropui ooeltzkuwi *. 

Malagasy Eegion. 
Madagascar ; — 

Eidolon dupreannm *. 

Pteropus rufus* nifus (northern and central) ; riifiis priiiceps (southern). 

Comoro Islands : — Pterojms comorensis * ; liningdnnei *. 

Aldabra : — Ptcrojius aldabrensis *. 

Seychelles : — Pterojms seychettensis *. 

Mauritius, Reunion : — Pteropus suhiiger * ; niger * ; ? roilricensis. 

Rodriguez : — Pteroptis rodricensis * (perhaps also Mauritius). 

Indian and Ceylonese Subrkgioks. 

Maldives : — Pteropus ariel*. 

Ceylon : — 

Rousettus seminudus *. 

Pteropus giganteus giga7}teiis (also Indian Peninsula). 

Cynopterus sphinx sphinx (also Indian Peninsula); brachyofis ceylonensis*. 

Indian Peninsula : — 

Rousettus arabicus (Karachi ; also S. Arabia) ; leschenaulti (also Indo- 

Pteropus gigantetis giganteus (also Ceylsn). 
Cynopterus sphinx sphinx {^. &E. provinces; also Ceylon, Assam, N. Burma, 

N. Siam) ; sphinx gangeticus (N.W. & C. Provinces). 

Indo-Chinese Scbregion. 

Himalayas (Kooloo, Nepal, Darjeeling, Assam, Cachar, Manipur) : — 
Rousettus leschenaulti (also Indian Peninsula, Burma, &c.). 
Pteropus giganteus leucocephalus *. 
Cynopterus sphinx sphinx (Assam ; also Indian Peninsula, N. Buraia, 

N. Siam) ; brachyotis angulatus (also Burma, Siam, &c.). 
'i Macroglossm minimus sobrinus ("Darjeeling"; also Malay Peninsula, 

&c. ; perhaps Siam, Burma, Tenasserim). 

Burma : — 

Rousettus lescheimidti (also Indian Peninsula, Siam, &c.). 

Cyno'pterus sphinx sj>hinx (N.Burma; also Indian Peninsula, N. Siam); 

brachyotis angulatus (also Assam, Siam, &c.). 
Spharias * blanfordi. 

Eonycieris spelaa (also Tenasserim, Siam, Malay Peninsula, <fec.). 
? Macroglossus minimus sobrinus (also Malay Peninsula, &c. ; perhaps 

Siam, Tenasserim). 

Tenasserim : — 

Pteropus intermedins {*) (probably also Lower Siam). 
Eonycieris spelcea (also Burma, Siam, Malay Peninsula, &c.). 
? Macroglossus minimus sobrinus (also Malay Peninsula, Sumatra, &c. 
perhaps Burma, Siam). ,y..,.,,. ,-._....,.■.. 


Mergui Archipelago ; — Ptcropus ki/pomelanus geMinorum *. 

South China (Amoy) : — Boiisettus Icschenaulti (also Siain, Burma, Nepal, 
Indian Peninsula). 

South Liu-kiu Islands : — Piernpus dasymalhts * ; loochoensis *. 

Formosa : — Pteropus formosics *. 

Siani, Cambodja, Saigon, Pulo Condor: — 

Buuscf/us ksvhenaulti (Slam ; also S. China, Burma, Nepal, Inijiau Penin- 
sula) ; ainplcccicaudiitus (Cambodja ; also Indo-Malaya). 

Pteropus hypomelamis condorensis * (Siam, P. Condor) ; It/lei* . 

Ci/nopteruA sphinx sphinx (also N. Burma, Indian Peninsula) ; hrachyofis 
angulatus (also Burma, Assam, Malay Peninsula, &c.). 

Eonyctcris spelcea (also Burma, Teuasseriw, Malay Peninsula, &c.). 

? Macroglossus minimus sobrinus (also Malay Peninsula ; perhaps Buiuia, 
Tenasserim, Darjeeling). 


Andamaus : — 

Pteropus safyrns* (Naroondam); tytleri*. 
Cynoptcrus brachyotis irruchysoma *. 

Kicobars: — 

Pteropus faunulus * ; melanotus *. 
Cynopterus brachyotis scherzeri *. 

Lower Siam : — 

1 Bouset/us amplexicaudafus (Indo-Malaya generally, Cambodja). 

? Pteropus intermedius (also Tenasserim). 

Cynopterus brachyotis angulatus (also Siam, Burma, Malay Peninsula, 

&c.) ; brachyotis brachyotis (also Malay Peninsula, Borneo, Sumatra, 

Eonycteris speltea (also Siam, Burma, Malay Peninsula, itc). 
1 Macroglossus minimus sobrinus (also Malay Peninsula, perhaps Siam). 

Malay Peninsula : — 

t Eousettus amplexicaudatvs (Indo-Malaya generally, Cambodja). 
Ptcropus va/iipyrus malaccennis (also Sumatra). 

Cynopterus brachyotis angulatus (also Lower Siam, Borneo, Sumatra, &c.) ; 
brachyotis brachyotis (also Lower Siam, Borneo, Sumatra, &c.) ; harpax *. 
Megarops ecaudatus (also Sumatra, Borneo). 
Penthetor lucasi (also Ehio Archipelago, Borneo). 
Eonycteris spelcea (also Siam, Burma, Sumatra, &c.). 

Macroglossus minimus minimus (?) (also Java, perhaps Sumatra); minimus 
sobriniis (also Sumatra and Java; perhaps Lower Siam, &c.). 

Sembilan Islands (off west coast of Malay Peninsula) : — Pteropus hypomelanus 
robinsoni *. 

Sumatra (incl. Rhio-Linga Archipelago. Banka): — 

h'ouseltus amplcxicaudatus (Indo-Malaya generally). 

Pteropus vampyrus malaccinsis (also IMulay Peninsula). 

Cyivrpterus sphinx titthcecheilus (also Java, Lombok) ; Irrachyotis angulatus 

(also Malay Peninsula, Sinialu, Mentawei) ; brachyotis brachyotis (also 

Malay Peninsula, Borneo, &e.); horsfieldi lyoni*. 
Mcgierops ecaudatus (also Malay Peninsula, Borneo). 
Penthetor lucasi (Khio Archipelago, so far not recorded from Sumatra ; 

also in Malay Peninsula, Borneo). 
Eonycteris spelcea- (also Malay Peninsula, Siam, Java, &c.). 
Macroglossus minimus minimus (?) (also Java, perhaps Malay Peninsula) ; 

minimus sobrinus (also Malay Peninsula, Java). 

Simalu Islands ; — Cynopterus brachyotis angulatus (also Mentawei, Sumatra, 


Nias ;— 

Pteropiig iriddieics *. 

Ci/!iopf(ri(s hruchyo is miiinlus * ; major * ; princejK *. 

MacnH/loasus minimus sobrinus (also yuniatra, Java). 

Mentawei Iftlixncls : — ■ 

? Plerapiis ht/pomehnics oiffwius (;ilso Engatio). 

Cyiiopfcrus hrachi/u/is anyidaius (also 8iiiialu, Sumatra, &c.). 

Engano : — 

Bousei/us amplexicttiidafiis (Indo-M:ilaya geiTerallr). 

Pierojms fiypomelaiius eiiyanns (*) (pei-liii)i3 also Mentawei) ; modiz/Hcrnii *. 

C'lii'istnuis Island (S. of Java) ; — P/cropus yiatalis*. 

Norib Nattmas : — 

Pferopns hypcmelunus cnniix * ; reinipi/riis natunee (also Borneo). 
? Cynoptcriis brackyotis brachyotis (ulso Malay Peninsula, Borneo, Sumatra, 

South Natimas ; — Pteropus hypomelanics annccfcns *. [Pteropus vampynis 
natuucs, not yet recorded.] 

Tambelan Islands, Pnlo Aor, Tiomari, Lantingii, Great Redang, Perbentiau : — 
Ptcropns hypomelaiiits lepidus*. 

Borneo (incl. Lnbiian, Mengalim, Sibufii, Lamboyan Islands) : — 

Jiouseiins amjilcricandafus (Cambodja, Indo- Malaya generally)- 

Pteropus hypomelamis fomesi* ; speciosus (Sibutu I. ; also Malanipa I., ofT 
Zamboanga) ; vampyrus naiuniE (also N. Natunas). 

Cynopta-us brnchyotis braehyotis (also Malay Peninsula, Sumatra, Philip- 
pines, CelKbes). 

HJeqcsrops efav.datns (also Malay Peninsula, Sumatra). 

ByacirphTits * spcidicens. 

llalionycti:ris * lunculaia. 

Peiithefor lucid (also Malay Peninsula, Rliio Archipelago). 

Eonycteris major *. 

MacroyloSfJis higvchilus lagochilus (also Philippines, Celebes, &e.). 

Mata Siri l.slands : — Cynapienis bracliyoiis ivsuhtruiii (also ICangean Islands). 

Java (inol. Madura) : — 

lioitscitiis aJwrtridgei * ; minor *. 

Pleropus vampyriis vampyrus*. 

Cyiiopierus sphinx titihcechciius (also Sumatra, Louibok) ; brachyo/is 

javaniciis * ; hori^fieldi liorsficldi *. 
Chironax * mdauocephali(s. 

Eonycteris speUea (also Sumatra, Malay Peninsula, Siam, Burmn). 
31acrog/,o<'sus miHimus minimus {nhn Kangean Is.; perhaps Sumatra and 

Malay Peninsula); minimus sobrimts (also Sumatra, Malay Peiiiusula ; 

perhaps Indo-China). 

Eawean Islands: — Ptcropiis atcrrivms (also Kangean Is.). 

Kangean Islands :^ 

Pteropus aierrimus (also Bawean Is.). 

Cyuoptertis brachyotis insulariim ^als(> Mata Siri Is.). 

Macroglosstis minitnus minimus (also Java, &c.). 

B;ili : — Pteropm vampyrus pi utnn (also Lombok). 

Piiilippines (incl. Cagayan Sulu and Palmas Is.): — 

Jioiiseiius amphxicaudaius (Indo-Malaya generally), 

Pteropus hypomelanus caqai/avvs * ; sp?ciosus (Malanipa I., off Zam- 
boanga ; also Sibutu I., off N. Borneo); w/?««s (also Celebes) ; puiuilus* 
(Palmas Is.); leucopferus* (Luzon); vampyrus lancnsis* . 
'' Acerodoii jubotus* jubatus (Luzon, Leyt^, Negros, Dlnagat, ? Panay) ; 
jubatus mindanensis {'M\\\i\a\VM-); Incifrr* (Paimy). 

GF.ofiRArnicAi, DrsTRinuxroN. Ixxi 

Hai-pf/ioiii/ctci-L'i * whiteheadi (Jlindanao) 

Ci/nopterus braahi/ofis hrackyolis (also Borneo, Celebes, kc). 

J fci.'ocliirus *jnguri. 

Macrof/lossus lugochiliis (also Borneo, Celebes, &c.). 

,, , , AuSTKO-MaI.AVAN SuilKEGlON. 

( elebcs : — 

Riiusettits celebensis (also Sangliir). 

Buiieia * hidens. 

Pteroptis hypomelanua i)iaca.i$nricus (also Sanghir, Talaut) ; huiiihs (also 

Piiilippines) ; doiisuni'^ ; pfCAW/rf/z/s (also Gilolo and Amboina groups) ; 

alccto (also Salayer. LomSok). 
Accrodon cclehcnsis (also ijalayer, Sula Is.). 
Styloctcniam * w lUarci. 
Dolisonia cxolcta *. 

Ci/nopterus hrachyoih hrachynlis (also Borneo, riiilippiiies, kc). 
TliooptiTiis iiifjresccns (also Gilolo group). 
yyctimetie minutus* ; cephaloles (also Auiboina group, Timor Laut, 

fhnycteris rosenhcrgi *. 
Macrog/iissiiJi lagochilits lagochilus (also Borneo, riiilippines, Sanghir, 

Aniboina group). 

Sangliir Islands : — 

Ji'ousefti/s celchensis (also Celebes). 

r/rropiis /i.ypome/anit.f macassaricus (also Celebes, Talaut) ; rankeps (also 

Gilolo group, Sula Is.); ? mdanopttgon (also Amboina group, Banda Is., 

Timor Laut) ; ? chrysoprocius (also Amboina group). 
Macroghfsits lagochilus lagocliUus (also Celebes, Philippines, Avnboina 

group, &c.). 

Talaut Islands (Lirong) : — 

P/eropiis hypomclanus macassar tens (also Sanghii-, Celebes). 
Accrodon htimilis *. 

Bonerato, Dyampea, Salayer : — 

I'Icropiui griscus (also Timor) ; alecfo (also Celebes, Lombok). 
Accrodon celchensis (also Celebes, Sula Is.). 

liOmbok : — 

Picrnpits Inmhoccnsis (also Flores); vampi/rus pfiilon (also Bali); nlccfo 

(also Salayer, Celebes). 
Cynopterus sphinx titthmcheilitii (also Java. Sumatra). 

Flores : — 

Kouseltus amplexieauddfus (Indo-Malaya generally). 

P/eropiis lomhocensis (also Lombok). 

AcerodoH iiirm/clo/ijlore/sii *. 

Dohsoniu pcroiti (also Timor, Alor, Wetter). 

Sumba : — 

Ptcropits morio (also SavuJi 
Accrodon gilrus *. 
Dijhsonia ssuuihana *. 

Savn : — 

Rouftttuf. amplcxicaiidatiis (also Flores, Alor, Timor, Inilo-AIalaya). 
Pteropiis vampyrus edulis (also Timor) ; morio (also Siuuba). 

Timor (incl. Wetter): — 

Jiouscttits amplcxicaudutus (also Flores, Saru, Alor, Iiido-AIalaya^. 
Picropux gri.-tciis (also Bonerato, Dyampea); tenimiihcki (also Amboina 

group) ; vampyrns edulis (also Savu). 
Acrodon mnckloli inacklofi *. 

Djbsonia moluccensis (?) (also Amboina group, Aru Is.) ; pcroni (also Flores , 


? Cyiioptenis sphinx tiiihacheilus (also Lombok, Jaxa, Sumatra). 
Nyctime'iie cephaloies (also Celebes, Aiuboina group, Timor Laut). 

Alor (Ombay) : — 

Eousettus amplexicaudatus (also Flores, Savu, Timor, Indo-Malaya). 

Pieropus soli/arius *. 

Acerodon mackloti alorensis *. 

Dobsonia peroni (also Flores, Wetter, Timor). 

Gilolo group (Morotai, Gilolo, Ternate, Batchian): — 

Pieropus hyponielanus hypomelanus * ; caniceps (also Snla Is., Sangliir) ; 

persmuitus (also Celebes, Amboiua group) ; chiysauchen (also Gliebi, 

Salawati, Mysol, N.W. New Guinea). 
Dobsonia crentdata *. 
2'hooptenis nigrescens (also Celebes). 
Nyciimene albiventer *. 

Sula Islands : — 

Pieropus canice-ps (also Gilolo group, Sangliir). 
Acerodon celebensia (also Celebes, Salayer). 

Amboina group (Buru, Amboina, Cerani, Goram, Manawolba, Watubella) : — 
Eouseiius hrachyotis (also New Guinea, &o.). 
Pieropus Hops*; "i argentaius* ; melanopogon (also Banda Is., Timor 

Laut, probably Sanghir) ; chrysoproctus {*) (perhaps also Sanghir) ; 
- temmincJci (also Tiuior) ; personatus (also Celebes, Gilolo group) ; 

ocularis *. 
Dobsonia moluccensis (also Aru Is., perLaps Timor); viridis umbrosa (also 

Banda Is.). 
Nyciime^ie varius * ; cephafotes (also Celebes, Timor Laut, Timor). 
Macroglossus lagochihis lagocJiilus (also Banda Is., Celebes, Philippines, 

Syconycteris crassa major *, 

Banda Islands : — 

Pieropus pallidus * ; melanopogon (also Amboina group, Timor Laut, 

probably Sanghir). 
Dobsonia viridis umbrosa (also Amboina group). 

Macroglossus lagochilus logochilus (also Amboina group, Sanghir, Celebes, 

Timor Laut: — 

Pieropus melanopogon (also Amboina group, Banda Is., &c.). 
Kyctimetie cephaloies (also Timor, Amboina group, Celebes). 

Key Islands ; — 

Pieropus keyensis *. 

Dobsonia viridis viridis *. [D. moluccensis not recorded, but possibly 

Occurring ; known from Amboiua group and Aru Is.] 
Nyciimene pajmanus (also New Guiuea, &c.). 
[Macroglossus lagochihis names not recorded, but possibly occurring ; 

known from Aru Is., New Guinea, &c.] 
Syconycteris crassa keyensis *. 

Aru Islands : — 

Pieropus aruensis * ; wacroiis *. 
Dobsonia moluccensis (also Amboina group). 
Macroglossus lagochilus nanus (also New Guinea, &c.). 
Syconycieris crassa papuana (also New Guinea). 

Ghebi, Salawati, Mysol, Waigeou : — 

Piei-opus chrysnuchen (also Gilolo group, N.W. New Guipea) ; 1 papuunus 

(also New Guinea). 
Dobsonia magna (also New Guinea). 
Nyciimene a'ello (also New Guinea). 
Macroglossus lagochilus nanus (also New Guinea, &c.). 


New Guinea : — 

Bousettus bra<;hyofis (also Amboina group, Bismarck Arch., Solomon Is.). 
Ptero/ms hi/pomekinus luteus (also islands off S.E. New Guinea) ; chrys- 

auchen (N.W. New Guinea; also Mysol, Salawati, Ghebi, Gilolo group); 

conapicilkifus (S.E. New Guinea with satellite islands, N. Australia) ; 

papuanus(*) (perhaps also Ghebi, Mvsol) ; epularius*. 
Dohsonia minor* ; magna (also My.sol, Waigeou). 
Nt/ctimene papuanus (also Admiralty Is., Key Is., Cape Yort) ; cyclotis * 

(Arfak Mts.) ; ccrfans* ; yeminus (also Trobriand and D'Entrecasteaux 

Is.) ; aello (also Mysol). 
Macroglossns lagochilus nanus (also Mysol, Bismarck Arch., Aru Is.). 
Syeonycteris crassa papuana (also Aru Is.). 
Alelonycieris melanops (also Bismarck Arch.). 

Conflict, Trobriand, D'Entrecasteaux, Alcester Is., Louisiades: — 

Pieropus kypomclainis luteus (also New Guinea) ; conspicillaius (also 

S.E. New Guinea, N. Australia); ? poliocephalus (also Australia). 
Dobsonia panuietensis *. 
Nyctinicne gemmus (also New Guinea). 
Syeonycteris crassa crassa *. 

Woodlark Island ; — 

Pteropus hypomelamis Inteus (also Trobriand and Conflict Is., New 
Guinea) ; conspicillatus (also Alcester and Trobriand Is., New Guinea, 

N. Australia). 
Nyctimene lullulce *. 
Syeonycteris naias *. 

Murray Islands (Torres Straits) : — Macroglossns lagochilus pygmaus*. 

Admiralty Islands : — 

Pteropus admiralitatum *. 

Nyctimcne papuanus (also New Guinea, &c.). 

Bismarck Archipelago : — 

Botisettus hrachyotis (also New Guinea, &c.). 

Pteropus capistrafus * ; neoliihernicus* . 

Dobsonia pradatrix ■*. 

Nyctimene major *. 

Macroglossus lagochilus nanus (also New Guinea, &c.). 

Syeonycteris crassa finschi *. 

Melonycteris melanwps (also New Guinea). 

Solomon Islands (whole Megachiroptera fauna) : — 

Bousettus brachyotis (also Guadaloanar, Bismarck Arcb., New Guinea). 
Ptercrpus colmius * ; solomonis * : cognafus * ; rayneri * ; ruhianus * ; 

lavellanus * ; grandis * ; ivoodfordi *. 
Pteralopex * anceps ; atrata. 
Dohsonia. inermis * ; nesea *. 
Nyctimene scitulus *. 
Macroglossns lagochilus yniavtus*. 
Nesonycteris * woodfordi. 

Known distribution in the separate groups of Soloiuon Islands; — 

Bougainrille, Shortland, Fauro : — 

Bousettus brachyotis (also Guadalcanar). 

Pteropus coloiius * ; grandis*. 

pteralopex aiiceps*. 

Dohsonia n^sea (also Eubiana). 

Nyctimene scitulus (also New Georgia, Florida, Guadalcanar). 

Nesonycteris woodfordi (also Guadalcanar). 

Vella I^vella : — Pteropus lavellanus*. 


Gliizo, Riibiana, New Georgia : — 

Fteropus iolomoHis* ; rubianus * ; woorfjunh (also Giiachilcaiiar). 

Bohsunia iiesea (also Sliortlaiid). 

Nyctimene scitulus (also Sliortlancl, Florida, Giiadalcauar). 

Florida, Giiadalcauar: — 

Emisettits hmchyotis (also Fauro). 

I'teropus rayneri * ; uoodfordi (also New Georgia). 

I'teralopex atrata *. 

yyctimene scitulus (also Shortland, New Georgia). 

Macroglossus layachilus microtiis '^. 

Nesonycteris woodfurdi (also 8horlland, Fauro). 

San Christoyal, Ugi : — 
Fteropus coynutus *. 
iJohsonia incrmis*. 

Australian Continent. 

ricrajiiis briinncns* ; youldi* ; conspiciUatus (also S.E. New Guinea and 
adjacent islands) ; i)olwccplialus (*) (perhaps also Trobriand Is.) ; 
scapula! us *. 
JS'yciimene pctpuanus (also New Guinea, &(;.) ; robbtsoni *. 
,»7' t<)jconijctcrU australis*. 

Polynesian Subregios. 

Bonin and Volcano Islands : — Fteropus pselaphon *. 

Mariannes ; — Fteropus mariaunus*. 

I'elew Islands : — Fteropus pelcwcnsis * ; pilosus *. 

Carolines (all islands togetlier) : — 

Fteropus yapeiisis * ; ualanns * ; moloss/nus * ; insnlaris * ; ph<Tcccphaliu> *. 
1 Nutoptcris macdomddl (also New Hebrides and Fijis). 

Records from the separate islands : — 
Yap, Mackenzie : — Fteropus yapensis *. 
Kuck Atoll : — Fteropus insularis *. 

Mortlock : — Fteropus molossinus (also Ponape) ; pluEoccplialua *. 
Ponape : — Fteropus molossinus (also Mortlock). 
TJalan (Kushai) : — Fteropus ualemus *. 

Sta. Cruz Islands (Vanikoro) : — Fteropus vanikoreusis * : ? tuherculatus * (there 
is Some uncertainty about the liabitat of this species). 

New Caledonia; — 

Fteropus ornatus * ; ycddici (also New Hebrides). 
Xotojjtcris neocaledonica *. 

Loyalty Islands : — Fteropus auratus *. 

New Hebrides (Aneiteuni) : — 

Fteropus ycddici (also New Caledonia) ; anetiauui^*. 
Notoptcris macdonaldi (also Fijis). 

Fijis: — 

Fteropus tovyanus (also Tontras, Sanioas) ; nanairusis *. 
Notoptcris macdonaldi (also New Hebrides). 

Tongas ; — Fteropus ioiiyanus (also Fijis, Sanioas). 

Saiuoas; — Ffcmpus ionyaiuis (also Fijis, Tongai-) ; famohiMS*. 


3. Jiemarls on the geoi/rcqjlucal distrihation. 

The northern limits of the area known to be inhabited by Mega- 
chiroptcra are Seneg:ambia, Egypt, Cyprus, Syria, South Arabia, 
Karachi (one unverified record of I'teropus girjanteus from Kelat), 
Himalayas, Amoy, Formosa, South Liu-kiu Islands, Bonins, Pelew 
Islands, and Mariannes ; in I'olynesia they occur as far east as the 
Samoa Islands. No Pruit-bat is known from North Africa west 
of Egypt and north of Senegal, nor from Asia Minor, Persia, Japan 
proper (records in literature of Pteropus dosi/mallus from Kiushiu 
and Hondo probably refer to captive specimens), Hawaii, New 
Zealand, and Tasmania (one ])robahly erroneous record of riero/ms 
poJiocephalus). The only fossil Fruit-bat thus far described is 
Archcopteroptis irmisiens, from the Upper Oligocene of Italy 
C' llousettus" (/aillardi, from the Middle Miocene of France, was, 
judging from 'the published figure of its humerus, a species of 

Of the four primary sections of Jfegachiroptera, one, the Epomo- 
]i7iorHS section (8 genera, 17 species, 19 forms), is strictly con- 
fined to the Ethiopian region ; a second, the Cjinopterus sectiou 
(11 genera, 31 species, 41 forms), extends from India and Ceylon 
in the west to the Solomon Islands and Australia in the east, and is 
represented bv one genus {Mijonyctevls, -4 species) in West Africa ; 
a third, the Sfacroqlossina' (7 genera, 13 species, iil forms), ranges 
from'Indo-China eastward to the Fiji Islands, and reoccurs, in a 
single genus and species (Mer/ahr/lossvs), in West Africa ; the 
fourth, the liouseltus section (9 genera, V2o species, 1-17 forms), in 
the number of species more than twice as large as the three other 
sections together, covers the whole of the area inhabited by Mega- 
chiroptera, from West Africa east to the Samoa Islands. 

A few genera have a very wide distribution : Ihnsettits (l-i species) 
from West Africa to the Solomon Islands (but not to Australia), 
Flernpus (So species, 103 forms, the largest of all genera, in the 
number of species nearly equal to all the other genera together) 
from the island of Pemba (south of Zanzibar), through the Mala- 
gasy, Oriental, and Australian regions to the Samoa Islands; 
others have a moderately wide range, e. g. Macroghssus from Indo- 
China to the Solomon Islands (not to Australia), Chinopteras from 
India and Ceylon to Celebes. But for the mnjority of genera the 
geographical "limits are much narrower, and not a few (chiefly 
monotypic genera) are so far known only from one place, island, 
or group of islands : Spharias from Burma only, Dtiacopterus and 
Balioni/clfiris from Borneo, Chironax from Java, Uarpi/ioni/cteris 
and Pienochints from the Philippines, Boncia and ^Slglocieniiim from 
Celebes, Pteralopex and Nesoivjrteris from the Solomon Islands : all 
I'lpomophorine genera, c:s.ce\>th: porno phorus, as well as Mi/onifderis 
and Megnhglossvs are practically confined to the whole or part of 
the great West African Forest Tract. 

The range of the species is, of course, as a rule restricted within 
much narrower limits, or if a species is widely distributed it has 
usuallv difl'erentiated into a number of local forms; Pteiopvs hi/po- 


vtelavus, for instance, covers practically the whole area from Indo- 
China to New Guinea and Woodlark Island, but is broken up into 
no less than twelve subspecies ; Fteropus vampyrus ranges over the 
whole of the Indo-Mala3an subregion, but divided into six races ; 
IJjiomoj^horvs wuhlbergi is distributed over the greater part of the 
Ethiopian region (except the Guinea coast west of Canieroons), but 
falls into two races — a western and eastern ; Cynopterus bmchyotis 
is known from tho whole of Indo-Malaya, extending eastward 
beyond this subregion to Celebes, westward to Kurma and Assam, 
and reappearing farther west in Ceylon, but is broken up into eight 
local forms ; MacroyJossus Ingocliilus extends from Borneo in an 
unbroken range eastward to the Solomon Islands, but has differ- 
entiated into four subspecies ; Syconycteris crassa ranges from tho 
Moluccas to the islands south-east of New Guinea, but has split 
into five races. There are, however, a few noticeable exceptions 
from the general rule of the relatively narrowly limited range of 
the species and subspecies; Eidolon hdinmi is generally distributed, 
without any appreciable change of characters, over nearly the 
•whole of the Ethiopian region (South Arabia excepted) ; Itousetius 
ampJexicaudatus occurs in Cambodja, through the Philippines, 
Uorneo, Sumatra (not Java), east to Flores and Timor, apparently 
without splitting into local forms ; Bousettus brctcliyotis ranges, 
seemingly unchanged, from the Amboina group to the Solomon 
Islands ; Cynopterus bracJiyotisbrachyotis from Lower Siam, through 
the Malay Peninsula to Sumatra, Eorneo, the Philippines, and 
Celebes ; Mncroylossus layochilus lagochilus over Borneo, the Philip- 
pines, Celebes, and the Amboina group. Whether these exceptions 
are really much more frequent and much more striking among 
Megachiroptera (and Chiroptera in general) than among non-flying 
Mammalia, is perhaps doubtful. In any case, against these few 
instances of an exceptionally wide distribution of one apparently 
unchanged form may be placed a large number of species known 
only from, and in many cases probably really restricted to, one 
island or group of islands. 

The evidence afibrded by the geographical distribution of Bats 
has generally been considered of doubtful value ; hence they have 
either been entirely excluded from the material worked out by 
zoogeographers or at least treated with pronounced suspicion, as 
likely to be more or less unreliable documents of evidence. This 
unwillingness or hesitation to place Bats on an equal zoogeographical 
footing with non-flying Mammalia would seem to be due, partly to 
the preconceived idea that owing to their power of flight Bats must 
evidently have been able easily to spread across barriers which, in 
ordinary circumstances, are insuperable for wingless Mammalia ; 
partljf to the fact that hitherto very often whole series of distinct 
forms have been concealed under one technical name. So long as 
(to mention only three cases among many) " Macroglosstis minimus" 
was believed to range unchanged from the Himalayas to New 
Guinea, Australia, and the Solomon Islands (now two distinct 
genera, thirteen recognizable forms), or " Cynoi-iUrvs margiiuilus " 


over India, Ceylon, Indo-China, and Indo-Malaya (now six species, 
fourteen forms), or " Rhhiolopfms ferrum-equiniim" uniformly over 
Europe, Asia, and Africa (now numerous distinct forms), they were 
undoubtedly of questionable value as zoogeographical material, 
liut these and similar anomalies invariably disappear as soon as 
modern methods of discrimination apijlied on vastly increased 
material render it possible to draw the lines of separation between 
the species (and their local modifications) somewhat more closely 
in accordance with the lines drawn by Nature. The second argu- 
ment referred to above, that the spreading of Bats from one locality 
to another must obviously have been greatly facilitated by their 
possession of wings, may in theory appear plausible enough, but 
when tested on the actual distribution of the species and subspecies 
it proves to be of much less importance than commonly supposed ; 
it rests, in reality, on a confusion of two different things : the 
j)Ower of flight no doubt would enable a Bat to spread over a much 
larger area than non-fiying Mammalia, but, as a matter of fact, 
only in very few cases is there any reason to believe that it has 
caused it to do so. The following pages, in which the distribution 
of the Alegachiroptera within each zoogeographical region or sub- 
region is discussed in some detail, will give ample evidence to this 
effect, but a few of the more striking examples may be mentioned 
here : a species of Pleroj)us inhabits the island of Pemba, south of 
Zanzibar, but although this island is separated from Africa by 
a channel only o5-40 miles wide, not this particular species only 
but the whole genus is unknown from any part of the adjacent 
continent ; although absent from Africa the genus Pteropus is 
distributed all over the Malagasy region, and each group of 
islands (Madagascar, Comoros, Aldabra, Seychelles, Mascarenes) has 
its own peculiar species, intermigration between the groups of 
islands is unknown ; the Epomophorine section of Fruit-bats is 
distributed over the whole of the Ethiopian region (eight genera, 
nineteen forms), but not a single form has spread to any island of 
the Malagasy region ; the Fteropus melanotiis group of species 
is distributed over the Andamaus, Nicobars, Nias, Engano, and 
Christmas Island (south of Java), and the whole group is confined 
to this chain of islands, no form having spread to the neighbouring 
Malay Peninsula or Sumatra; Fteropus hi/pomeJanus is represented 
by a local form in Engano, off Western Sumatra, but the species, 
though widely distributed elsewhere in Indo-Malaya, is unknown 
in Sumatra and Java ; the Megachiropteran fauna of Ceylon shows, 
of course, very close affinities to that of the Indian Peninsula, but 
the Indian llouseittis lesclienanlti is replaced by a distinct species 
{seminudus), and the Indo-Chinese and Indo-Malayan Cynopterus 
hrachi/otis is represcntea by a local form, though the species is 
unknown in the Peninsula ; the Eruit-bat faunas of the Malay 
Peninsula, Sumatra, and Borneo are closely interrelated, like their 
Mammalian faunas in general, but each has some distinct autoch- 
thonous forms of Eruit-bats (Borneo even two autochthonous 
genera), as it has of other ifammalia ; the Javan Mammalian fauna in 


general is more peculiar, both by the absence of some of the forms 
found in Sumatra and the Malay Peninsula and by the greater 
percentage of autochthonous forms, and this is again borne out by 
the Megachiropteran fauna of the island (absence of two Sumutran 
genera, presence of one autochthonous genus, among ten forms six 
autochthonous); the Ftero^nts rui/neri group is represented probably 
all over the Solomon Islands, but it has ditferentiated into five 
distinct species, one in the Bougainville group, a second on Yella 
].avella, a third in the New Georgia group, a fourth on Guadal- 
canar, and the fifth on San Christoval. This (and a series of 
similar examples might easily be adduced) tends to show that the 
present distribution of the Megachiroptera has not been influenced 
to any great, and as a rule not even to any appreciable, extent by 
their power of flight ; if it had, the Fruit-bat fauna of one group 
of islands could not, so commonly as is actnally the case, difl'er 
from that of a neighbouring group or continent, and the tendency 
to differentiation of insular species or forms would have been 
neutralized by the free intercourse between neighbouring faunas. 

Etldojiian Herjion. 

13 genera, 32 si^ecies (3-1: forms). Of the four primary sec- 
tions of Megachiroptera, one, the Bpomopltorus section (8 genera, 
17 species, 19 forms), is entirely Ethiopian ; of the liousettus 
section (9 genera, 125 species, 147 forms) three genera are repre- 
sented in this region, Eidolon by two species, liousettus by seven, 
and Pteropus by one ; the Cr/nopterus section (11 genera, 31 species, 
41 forms) by one peculiar genus, Myonycle)-is (four species) ; the 
Macroglossince (7 genera, 13 species, 21 forms) by one peculiar 
genus, Megalo'ihssus (one species). Of the thirteen genera ten 
are peculiar, viz. all except Eidolon (outside the Ethiopian region 
occurring only in Madagascar), llmis^ttus, and Fterojms. Of the 
tliirty-two species thirty are peculiar, viz. all except Rm'settus 
cef/ifptiacus, which extends to the eastern Mediterranean subregion, 
and liousettus arahiciis, which extends to Karachi. 

A subdivision of the Ethiopian region, on the basis of its 
Megachiropt«ran fauna, is very simple indeed. The region falls 
decidedly into two well-defined ])rovinces, viz. (1) the Wtst African 
Province, i. e. the whole of the Guinea coast and Congo basin, east 
to the western bank of Victoria Nyanza, south to Benguela and 
Damaraland, thus approximately synonymous with the Great West 
.\frican Forest llegion, though at least in the south-west extending 
beyond the limits of the forest belt, and (2) the East African 
Province, i. e. the eastern side of the continent from Erythrea and 
Abyssinia in the north to the Cnpc Colony in the south. So sharp 
is the contrast between the faunas of these two provinces that the 
large majority of genera and species occurring in the former are 
unknown in the latter, only a few species extending beyond its 
limits to part of East Africa. A single species, Eidolon Jiehnnn, 
seems to be thoroughlv common to both, ranging from Senegambia, 


Seminar, and Somaliluiid in the north, to Namaqualanrl, Mashona, 
and Nyosalaud in the south. Besides these two provinces, .South 
Arabia and the island of Peinba re(inire special remarks. — Matsehie's 
attempt to subdivide the Ethiopian region into a large number of 
zoogeographical districts ("Gebiete"), each with its own distinct 
Mammalian fauna, breaks completely down when tested on the 
distribution of the Ethiopian Fruit-bats. 

^\^est African provi)ice.- — 12 genera, 23 species C24 forms). 
Pterojms is the only genus which, though represented in the 
Ethiopian region (the island of Pcraba), is absent from W. Africa. 
Of the twelve genera no less than nine are peculiar, viz. Flerotrs, 
Epomops, Ifi/psi(jnafhi(S, Micropteropus, Kaiioiii/ctens, Scofoni/cteris, 
and Casini/cteris (all of the Epnmciphorus section), Mi/o»>/cteris 
(Ci/noptertis section), and MegaJoglossiis {31'(crofjJossincr) ; expressed 
in other words : all genera of the Epomophorine section of Eruit- 
bats are contined to this province, except Ej)omophorus itself, which 
is common to W. and E. Africa, and this province (" Afro-Malaya," 
as it was, perhaps a little jokingly, called by the late Dr. 11. Bowdler 
Sharpe) is the only part of Africa in which the otherwise essentaallj' 
Oriental and Austro-Malayan Cynopterine and Macroglossino 
sections are represented. The three non-autochthonous genera are 
Eidolon, lloasettxs, and Epomopihorns. The genus liousettus falls 
into three subgenera, liouseUns, Slenon}jcteriK, and Lissonycteris ; 
of these the first is widely distributed in the Ethiopian and Oriental 
regions and Austro-Malaya, while the two latter are confined to 
the Ethiopian region ; Steaoiii/cterin is common to the West and East 
African Provinces (one species in each), Lissoni/cteris (two sjieries) 
is peculiar to West Africa. Of the twenty-four "West African 
forms no less than twenty-one are peculiar, {. e. all forms except 
Eidolon helvum (generally distributed in the Ethiopian region), 
lioiisettan cvijfipliacus (Eoanda and Congo to Egypt, Palestine, &c.), 
and Epomophorus (jamhiivms (Guinea coast to Sennaar and Abys- 
sinia); only one of the twenty -one peculiar forms, viz. Epomophorus 
wahlbergi haldemani, extends eastward a little beyond the borders 
of the province, into German and British East Africa. 

Suggestions as to a possible subdivision of the province would 
be ])remature ; in too many cases the range of the genera and 
species is only imperfectly known. Some species (Ili/psignathus 
moiistrosus, Micropteropus pusillus) have a continuous distributit)n 
from Gambia, along the Guinea coast, through the Congo ]5asin to 
Victoria Xyanza ; in other cases the area is divided between several 
species of one genus : Epomops hurttilofcri in Liberia and Sierra 
Leone, E. fritnq\teti from the Gold Coast, east to Victoria Xyanza, 
south to Loanda, E.dohsoni in Bongucla and Katanga; Mi/oni/ctcris 
leptodon in Liberia and Sierra Leone, J/. wroiKjhtoni in the \\'e]lc 
district of the Congo Basin, M. torquafa in the Lower Congo district; 
and Angola. 

Islands in Gnlf of Ginnfn. — The followin<T species have been 
recorded from the ishituls of Eernando Po, Principe. Sau Thome, 


and Annobon : — Fernando Po, Eidolon helvum, Hypsignathns mon- 
strostis, Scotom/cteris zenlceri : Principe, Eidolon helvum : San Thome, 
Eidolon helvum, Myoni/cteris hrachycephcda : Annobon, Eidolon 
helvum. Myonycteris brachycejdiala is known only from San 
Thome, and the sole representative of the subgenus Phyyeiis ; all 
the other three species are common to the islands and the adjacent 
coasts of the continent. 

East African province. — 3 genera {Eidolon, Rousettus, Epomo- 
pJiorus), 10 species. This province is incomparabl)' poorer than 
W. Africa, both in genera and species and in the number of really 
peculiar forms. None of the genera is autochthonous. Eidolon 
helvum, Rousettus ^gyptiacus, and Epomophorus gambianus are 
common to this and the foregoing province ; the other seven forms 
(two Rousettus, five Ejjomojihorus) are entirely East African. 

Rousettus leaclii ranges from the Cape Colony north to British 
East Africa, and is in Egypt and part of West Africa replaced by 
R. cegyptiacus ; Epomophorus wahlbergi ivahlbergi likewise from the 
Cape Colony to British East Africa, and is replaced in West Africa 
by E. IV. haldemani ; Epomophorus crypturus extends from Trans- 
vaal to Nyasaland, and is replaced northward, from Tanganyika to 
Erythrea, by E. anurus. It will be noticed that the southern forms 
have as a rule an unbroken distribution from the Cape Colony or 
Transvaal north to British East Africa or at least to Nyasaland. 

South Arabia. — 2 genera, 2 species. Affinities decidedly African, 
J^i(fo?o)isa^f»!M/rt (autochthonous) being closely related to the widely- 
distributed African E. helvum, Rousettus arabicus to the East 
African R. leachi. R. arabicus extends as far east as Karachi, 
N.W. India. 

Pemba Island. — Pi.emarkable by being inhabited by a distinct 
species of Pt/ropms, although the genus is absent from the whole 
continent of Africa. The Pemba species {Pt. voeltzkowi) is closely 
related to a Malagasy form (Pt. comorensis). 

Malagasy Region. 

2 genera, 9 species (10 forms: Eidolon one, Pterojnts nine); 
neither of the genera is confined to this region, but all the species 
are peculiar. The single species of Eidolon (E. dupreanum, closely 
related to the Ethiopian E. helvum) is apparently restricted to 
the island of Madagascar, while Pterop)us is distributed over the 
whole of the Malagasy region. The affinities of the Malagasy 
Pteropus fauna have been discussed elsewhere (pp. 79-80). 

Indian and Ceylonese subregions. 

3 genera, 7 species (8 forms), viz. Rousettus (three species), 
Pteropus (two species), Cynoptcrus (two species, three forms). 
Rousettus arabicus is a South Arabian species which only enters the 


north-west corner of the Indian Peninsula ; li. le.Juncndti occur. 
all over the Teninsnla as well as in Lulo-Gluna, but i« in Ceyloa 
rep aced by a very closely related peculiar fori, H. semJdus. 
27nL^M7 %^^" ^"^''" ""^ Himalayan representative of 

tHvi 1 df M 7^> ■'■ r"^'^''"' (^'■""' ^^^'''^^ it only dm-ers in 
trivial details; Peninsular specimens (subspecies ^fuanieus) are 
nidistinguishable from Ceylonese, but slightly different from 
ilimalayau (subspecies leucocej^halns) a.d :\raldive specimens (««,?) 
Cpioptcrus spJnu.v has in continental South Asia differentiated into 
two subspecies, the one (sphin.v) ranging over the southern ar-d 
eastern provinces of the Indian Peninsula! south to Ceylon orth- 
east nortiern Indo-China, the other („eticus) known ojy 
rom the central and north-western provinces of India. In addition 
ioJtoMus ssminudns Ceylon possesses a second peculiar Pruit- 
bat, Cynopten.s brachi/olis ceylonensis, to which there is no 
equivalent in the Indian Peninsula, all the other forms of C. bracZ 
otis being Indo-ilalayan or Indo-Chinese. ^ 

lado- Chinese suhnyion. 
6 genera, 14 species (15 forms), viz. llonsetu,s (two species) 
l^eropus (seve.i species, eight forms), Cynopterus two ^"c e.) 
>Sp]ueruts (one), Eouycteris (one), Macroylossus (one). 

Continental Indo.China.-Roasettus JeschenauUi and Cynonteru, 
splnn. .pAuu- are invaders from west, the Indian Penmsuk 
h same may be said of the Himalayan Pteropus gigar.teus ulco- 
cei^alns, but it is subspecitically distinct from the Indian form 
Evidence of the affinities of the present area with Indo-ialaTa 
are, first the direct invaders from the latter; second, those forms 

closelj related to Iiido-ilalayan forms. To the first cate^orv 
beongi?o«.... a.n,ae.icaud[at.s, Eonycteris spekeJ^titZl 
rjlossus nunnnus. To the second, I'terojn^s inieJediusWenasserim) 
a representative of the Indo-Mala/an Ft. van>pyrL ■ S Si 
(Siam and Sa,gon), a peculiar species of the slme group and 
fi-hpomelanus condorensis (Siam, Cambodja, Palo Condor) a local 
race o a widey-distnbutcd Indo- and Austro-Malayan spedes 
Cynopterus brachyoUs anyulatus, rather than being an invader from 
wTrd'thr^::':^;'''^^ "' ^"^^-^^Z-- ^-^ -^ch h°as spread Luh- 
.fonh 8 a highly peculiar autochthonous genus and species of the 
Cynopterus group so far known only from Burma. 

Formosa.-T\x& single species recorded, Ptevopus formosvs, has 

ts closest relatives to the south in the Amboina groip (Pt. lops) 

to the north in the South Liu-kiu islands (Pt. dasymallus). ^ ^' 

Jl^^i/^-^^^-^-j^^i^lcmds.-Tbe two species of Pteropus known from 
these islands belong to different groups of the genus. Pt cZ 

ForL"an/y ,°^«"t-"-l -^--; ^'^ '^-^ repre^sentative of tte^. fonnosue; Pt. loochoensis a species of the otherwise 
entirely Polynesian Pt, rnariannus group. omerwise 


Ixxxii CEOGUArnicAL msTEiBuriorf. 

Indo-Malayan subrcgion . 

13 genera, JiG species (55 forms). Of the genera seven (all 
nionotypic) are peculiar, yiz. one of the Rousettus section [Dob- 
sonia liranch), Hn>-j)yionyrtens, and six of the Gijnoptervs section, 
Ptcnochiriis, Megteropx, Di/acopteru$, Balionycteris, Ch'rona.v, and 
Penthelor. Twenty-seven species (forty-six forms) are peculiar. 
Only one or two forms have entered the present subregion from the 
west (Indo-China), viz. Cijnnplerus sphinx (here differentiated as a 
distinct subspecies, titiJuec/ieiJvs) and, perhaps, Cynojitenis brachy- 
oiis angjilatus; but four species have spread in the opposite 
direction, from Indo-Malaj'a into Indo-China, viz. Rousettus amplexi- 
caudattis, Fieroptis hypomelanus, Eonycteris spelcva, and Macroglossus 
minimus. Macroylo^sus lar/ochilus is the only species which is 
widely distributed both in Indo- and Austro-Malaya, but four 
other Indo-Malayan species extend into the extreme western or 
south-western Austro-Malaya, viz. Pteropus mimus and Cynopferus 
brachyotis to Celebes, Rousettus amjjlcxicautlatus and Fierojms 
vampyrus to Timor. 

Malay Peninsula, Sumatra, and Borneo. — Taken as a whole this 
" province " is characterized, as against any other part of Indo- 
Malaj'a, by the presence of four peculiar genera, Megcerops 
(ecaudatits), Dyacoptenis {spadicevs), Balionycteris (mactdaia), and 
Pentheior (liicasi). It must be mentioned, however, that Balio- 
nycteris has its "equivalent" in Java {CJtirona.v), and Penthetor a 
close relative in western Austro-Malaya {Thoopterus). The status 
of the Malay Peninsula, Sumatra, and Borneo inter se is, judged by 
their Megachiroptera, practically the same as if judged by any 
other large group of Mammalia, •/. e. they are undoubtedlj' closely 
connected zoogeographically, while at the same time each has certain 
distinctive features, these latter decidedly more pronounced in the 
Eorneanthan in the Peninsular and Sumatran faunas. Borneo has 
two peculiar genera {Dyacnpterus, Balionycteris) and four peculiar 
forms, the Malay Peninsula and Sumatra have no peculiar genera, 
but each one peculiar form. The following forms arc common to 
all three areas, viz. Rousettus ample.cicaudatus, Cynop^terus brachy- 
otis brachyotis, Meya'rops ecaudatus, and Penthctor lucasi (not 
actually recorded from Sumatra itself, but from the E,hio Archi- 
pelago) : of these the two latter are the most important, since 
found nowhere outside this province ; the two former have a wider 
distribution in Indo-Malaya. The following are common to the 
Malay Peninsula and Sumatra, but unknown from Borneo, viz. 
Pteropus vampyrus malaccensis (in Borneo replaced by Pt. v. natuna:), 
Cynop)terus brachyotis angidatus, Eonycteris spelcea (in Borneo 
replaced by E. major), and Macroglossus minimus (in Borneo re- 
placed by a form of M. lagocMlus). No form is common to 
Sumatra and Borneo but absent in the Malay Peninsula, and none 
is common to the Malay Peninsula and Borneo but absent in 
Sumatra. distribution. Ixxxiii 

Andamans.~On\y Pteropus (t^ro species) and Ci,mpterus (a local 
form of C. hraclnjotis) l,avc found their way to these island lu 
hree orms are peculiar. Of the two s^Lies of Pter^^s, one 
(PL teller) belongs to the Pt. melanotus group, a group en irelv 
confined to the Andaman, Nicobar, Nias, Engano, and Chris Lv 
Ks and chain; the other (P. ...^..s-) to the Z 
lUufponrdanus group. C;faopten.s brachyotis is a common Indo- 
Chinese and Indo-l[alayan type. 

Nicohars.~L% in the Andaraans, one species of the Pteroom 
melanotas^ron^ i^Pt. ^nelanot.s), one of the Pt. k.ponulannZZ 
ifuaaidus) an^ one local form of Canopterm hraduioth, all throe 
torms peculiar, though closely related to those of the Andamans. 

Nias.—MacrngJossus minimus sohriaus (also in Sumatra^ • a 
J.ecuhar species of the Pteropus melanotus group (Pt nuulLJ 

rfhepf^^t'l '^ ''' ""''-'^^ ^^'^^'?^ ^' "^^^ I4^trt u : 

of the It. melanotus group is al,scnt in Sumatra); no less than 
three pecnliar forms of C.noj.terus, vi. one local «ubs^^ ies o 
C. hraIn,ot,s (a different subspecies in Sumatra), one species 
;h«/o.) allied to C. bracJnjotis anrpUatus (Indo-China Eu- 
loninsula, Sumatra), and one unusually well-differentiated snecies 

&r?;'j.t;.;^;sir""-"°° " "'■""""'• <-*--''^^" 

the widely-distributed Pousatus an,ple.ncaudalus. The differences 
from the fauna of the neighbouring Sumatra are strikit tha 
many Sumatran types are absent in tins outlying island, wa^ only 
o be expected; but the Pt. Jujponuhmus ^ud^melanous groups 
though represented m Engano, are unknown in Sumatra. ^ ' 

Christmas Island (S of Java).-One peculiar species oi Pteropus 
belonging to a group of the genus {Pt. melanotus) represented £ 
where only in Engano, Xias, the Alcobars, and Andamans. 

J-«,« The Fruit-bat fauna of Java, like ils Mammalian fauna 
n %nZk '' ^oma^-kable (1) by the absence of certain types foS 

Intorb , ^'°'"'"\ (') ^^' ^^' ''^^''''^^y 1-rge number of 

autochthonous forms, and (3) by its complicated affinities these 
poniting partly toward tlie neighbouring islands, part y w th 
exc usion of these latter toward Indo-ChinI To takJ thes^ thJee 
categories of faunistic characters separately :—(i) The followip<. 
iorms present both in Sumatra (and the Malav Peninsula) and 
Borneo are absent from Java, viz. Eousettus- ample.cicaJ(aC 
Cynopta-us hracujot.s hrach;,otis, Me.,a>rops (ccaudaL), and P.t' 
ITJT'^ . (2) Of ten forms known from Java six ar. pecu i 
one of these (C%u-o,m.. mdanocepludus) being a pecnliar genus 
(.5 a) Poruis affiuities with Sumatra, or Borneo, or boll 



viz. Bousettus 7ninoj\ the Javaii representative of B. 
eauJalus; Pieropus vnmpurus vojnpyrus, replaced in Sumatra by 
Ft. V. malaccensis, in Borneo by Ft. v. natunce ; Cj/nopterus hruchyotis 
javanicus, a local form of the widely-distributed C. b. hracliyotis ; 
Cunopterus horsjieldi horsjieldi, replaced in Sumatra by G. h. lyoni, 
while the species is absent in Borneo ; Chironax mclanocej)halus, 
represented in Borneo by BaJionycteris macidata, botli genera having 
no equivalent elsewhere; Eonyctcris speJcea, also in Sumatra, but 
in Borneo a distinct species {major) ; Macroghs^ns minimus, also in 
Sumatra, but in Borneo a distinct species (lagocTiilus) : (3 b) Evidence 
of affinities with Indo-China, with exclusion of Sumatra and Borneo, 
Bousettus shortridyei, closely allied to the Indo-Chinese B. lesche- 
naulti, a bat which has no rei)rescntative elsewhere in Indo-Malaya; 
probably also Cinmpterus spJiiny tittJuvcheihis (though this form is 
common to Java and Sumatra), Cynopterus sphin.v being decidedly 
not an Indo-ilalayan but an Indian and Indo-Chinese type, which, 
probably, in Java has ditferentiated into a distinct race, C. s. titthe- 
cheilus, which again has spread westward to Sumatra, eastward to 
Lombok, perhaps as far as Timor. 

Philippines. — 7 genera, 13 species (14 forms). The general 
characters of the Fruit-bat fauna (which is, of course, still im- 
perfectly known) would seem to be these : the autochthonous element 
is strongly developed (tv/o genera, one of which is so peculiarly 
modified that it has recently been proposed to separate it in a dis- 
tinct subfamily, six species, nine forms), the affinities of the autoch- 
thonous forms are partly Indo-Malayan, partly Austro-Malayan, 
while one species has its closest known relatives in the Bonin and 
JPolew Islands; in addition, some direct invasion has taken place 
both from Indo-Malaya (Borneo) and Austro-Malaya (Celebes), 
perhaps also, though if so to a much less extent, movements in the 
opposite direction. The details are: — Indigenous genera: Har- 
pyyionycteris (whiteheadi), a highly peculiar genus, with no nearer 
relative than the Austro-Malayan Dobsonia ; Ptenochirus {jwjori), 
closely allied to Cynopterus, and most closely to its " Niadius '"' 
section, which is known only from the Malay Peninsula, Sumatra, 
Nias, and Java. (2) Indigenous representatives of (essentially) 
Indo-Malayan species: Pteropus hypomelanus cagayanus; Pt. vam- 
pijnis lanensis. (3) Indigenous representatives of Austro-Malayan 
species: Pteropus pumdus, closely allied to Pt. griseus (Bonerato, 
Dyampea, Timor) ; Acerodon juhatus and lucifer, all other forms of 
Aeerodon being Austro-Malayan. (4) Indigenous representative of 
an otherwise exclusively north-west Polynesian group : Pteropus 
leucopterus, allied to Pt.pselaphon (Bonin Islands) and Pt. pelewensis 
(Pelew Islands). (5) Probable invaders from Indo-Malaya (Borneo) : 
Bousettus amplexicaudatus ; Pteropus speciosus (?) ; Cynopterus bra- 
cliyotis bracJiyotis ; Macroglossus lagochihis lagocldJus. (6) Invader 
from Celebes (or from the Philippines into Celebes) : Pteropus 


Austro-Mahtyati Suhrecjion. 

If) genera, 80 species (93 forms), thus much richer in species 
than any other subregion, the number of species being more than 
twice that of Indo-Malaya ; 43 per cent, of all known genera and 
exactly the same percentage of all known species are represented 
in this subregion. Of the fifteen genera seven (with altogether 
nineteen species) are peculiar, viz. four of the liousettus section, 
Boneia (one species), Ptcralopex (two), Sti/hctenium (one), and 
Dohsonia (twelve) ; one of the Ci/nopterus section, Thoopterus (one) ; 
and two Macroglossincp, Melonyctcris (one) and 2\esoni/ctens (one). 
The affinities of these seven autochthonous genera are as follow s : — 
Boneia is closely related to the widely-spread llousetius ; Fteralopeo' 
and Sti/Iocienium are offshoots from the even more widely- 
di.stributed Pteropns ; Dohsonia is an unusually peculiarly modified 
g( iius with only one near relative, the Philippine Harpyionycteris, 
and is both by its comparatively large number of species, all strictly 
confined to this subregion, by its many aberrant characters, and ])y 
its distribution over practically the whole area from Celebes to the 
Solomon Islands, zoogeographically perhaps the most important of 
the autochthonous genera ; Thoopterus is the Austro-Malayan repre- 
sentative of the Indo-Malayan Fentlietor x while Midonycteris and 
Nesonycteris have no closer relative than the Polynesian Notopieris. 
The non-autochthonous genera are. liousettus, Pteropus, Acerodon 
(found elsewhere only in the Philijipines), Cynoptents (only entering 
the extreme western part of Austro-Malaya), Nyctimene (entirely 
Austro-Malayan, except for one Australian species, and one common 
to New Guinea and Australia), Eonycteris (ranging from Indo- 
Malaya only to Celebes), Macroyhssus (from Indo-China to the 
Solomon Islands), and Syconycteris (entirely Austro-Malayan, 
except for one Australian species). If Nyctimene and Syconyctei-is 
were added to the number of autochthonous genera, and they could 
evidently be so without much error, the number of such genera 
would be no less than nine (out of fifteen), with altogether thirty- 
three species, so that the number of peculiar species belonging to 
peculiar genera would be 41 per cent, of the total number of 
species known from the subregion. 

Seventy-one of the eighty species (89 per cent.) are entirely- 
confined to Austro-Malaya. The nine species that are not wholly 
restricted to this subregion are : — (1) such as extend from Indo- 
Malaya only to the Lesser Sunda Islands, viz. Rousettus amplexi- 
cnudatus, Pteropus vampyr\is, and Cynopterus spJiin.v (subsp. titthce- 
cheilus) : (2) one extending from Indo-Malaya only to Celebes, 
Cynopterus bracJn/otis (suhsi^. brachyotis): (3) such as extend from 
Ihdo-ilalaya over a greater part of Austro-Malaya, but here differ- 
entiated into a number of subspecies, viz. Pteropus hypomelanus and 
Macroylossus layochilus : (4) one common only to the Philippines 
and Celebes, Pteropus mimus: (5) such as are common to Xew 
Guinea and Australia, viz. Pteropns ronspiicilJaius and Nyctimene 

ixxxvi geogra; nicAL w.siraisuTioN. 

Celebes and Sanr/Jiir Islands. — 11 genera, 17 species (excluding 
a few doubtful records from literature). Two genera, both mono- 
typic, are peculiar, Boneia, closely allied to liouscttus, and Stijlo- 
ctenium, closely allied to I'tero/nis and more particularly to the 
entirely Austro-Malayan I'L temminclci group, one species of Avhich 
{Ft. ]^iersoncUus) occurs also in Celebes. 80 far as the records go 
(for there is no doubt still much to be added to our knowledge of 
the Fruit-bats of Celebes and Austro-Malaya in general) seven 
species seem to be confined to Celebes, with the Sanghir islands, 
viz. liousettus celebensis, Boneia bidens, Pteropus dobsoni, Stylo- 
cteniiim wallacei, Dohsonia exoleta, Nyctimene minutus, and Eonyc- 
ieris rosenhergi ; and in connection with these may be mentioned 
Pteropus liypomelanus mucassnricus, Pteropus alecio, and Acerodon 
celebensis, though the first is known to extend north to the Talaut 
islands, the second south through Salaycr to Lombok, and the third 
south to Salayer and east to the Sula Islands. An aiiah'sis of the 
relationships of the fauna gives this result: — (1) Indigenous species 
pointing, geographically, in uncertain direction, liouseifus celebensis, 
a rather peculiar, narrow-toothed species of doubtful affinities, and 
Boneia bidens: (2) Indigenous species of an otherwise entirely 
Indo-Malayan genus, Eonycteris rosenberrji -. (3) Forms common to 
Celebes and some part of Indo-Malaya (Borneo and Philippines), 
Pteropus mi^nits, Cynopterus brachyotis brachyoiis, and Macroylossus 
layocMlus lar/ocJiilus : (4) Distinct form Avith quite close relatives 
both ill Indo- and Austro-Malaj'a, Pteropus I/ypomelanvs macas- 
saricus : (5) Distinctforms with clearly pronounced Austro-Malayan 
affinities, Pteropus dobsoni, Pt. alecto, Acerodon celebensis, Stylo- 
ctenium ivallacei, Dobsonia exoleta, and Nyctimene minutus : 
(6) Forms common to the Celebes and Moluccas (Gilolo group, or 
Amboina group, or both), Pteropus caniceps, Pt. personatns, Thoo- 
pterus niyresceiis, z^nd A'yciiinene ccplcalotes. The general conclusion 
is that, while the Indo-Malayan element is by no means incon- 
spicuous, the Austro-Malayan affinities of the Fruit-bat fauna are 
decidedlj' predominant. 

Amboina group (Buru, Amboina, Ceram, and smaller islands) and 
Banda Islands. — genera, 14 species. Of the Celebean genera, 
Boneia, Acerodon, Sfj/locfenium, Cynopterus, Thoopterus, and 
Eonycteris are absent, but Syconycteris has been added. No genus 
is autochthonous. Five species are (so far as known) restricted to 
this group of islands, viz. Pteropus pallidus (closely allied species 
in Celebes and Timor), Pt. Hops (no very close relative until in 
Formosa), Pt. cltrysoproctus (]K'rhaps also in Sanghir islands; 
curiously enough no closely related species until in the Solomon 
islands, where the gronj) is widely distributed), Pt. ocularis (closely 
allied species in the (iilolo group and New Guinea), and Nyctimene 
varius (a closely allied species in Celebes) ; to these may be added 
two peculiar subspecies, Dobsonia viridis umbrosa (another sub- 
species in the Key Islands) and Syconycteris crassa, major (other 
subspecies in the Papuan section of Austro-Malaya). The non- 

Gi:onK\ri[ic.\r. DisTr.iP.nioN. Isxxvii 

])Oculiar elemeui points towards any of the siirroundinf,' islands, 
the (jtikilo group. Celebes, South West Islands (Timor), .South East 
Islands (Timor Laut, Key, Aru), and New Guinea, it being in most 
cases mere conjecture whether the movements have taken place 
from the Amboina group into the neighbouring group or iu the 
o[)posite diicction. 

G'dolo ijroup (Morotai, Gilolo, Ternatc, Batjan, &c.). — 4 genera, 
7 species; but tlie fauna is no doubt more imperfectly known than 
tliat of the Amboina groiij). Tlie faunistic leanings are moro 
decidedly toward Celebes than to the Amboina grou[) and New 
Ciuinea, but a large number of Celtbean genera are absent from the 
list, viz. lloiisi'ttiis, Boneia, Acerodon, tSti/Ioctenium, C'l/nojiterns, 
Konijcteris, and Mcici'Of/lonsits; the absence of Iiousettus and Macro- 
ijlossiis, both generally distributed elsewhere in Indo- and Austro- 
Malaya, may be due to incompleteness of the I'ecords. As in the 
case of the Amboina group, no genus is autochthonous. Two species 
seem to be confined to the (iilolo group, Duhsonia cremtlata (a 
closely allied species in the Amboina group, viz. I), viridis) and 
jXiictimcne aJhiventcv (its closest relative iu New Guinea); in 
addition to tliese, one autochthonous subspecies of Pteropus hiqiO- 
rtnlamis (other subspecies in CelebcK and New Guinea, but not 
in the Amboina group). Specimens common with Celebes and 
Sanghir Islands are, Fieropus caniceps, Ft. 2>erso»a(us, and Thoo- 
jiierus nigresccns (genus confined to Celebes and the jjresent group). 
One species is common with N.W. New Guinea, Fieropus clirys- 

New Guinea (including Ghebi, Salawati, Mysol, and Waigeou, 
but excluding the satellite islands to the north, east, and south).— 
7 genera, lO species. The genera are tlie same as tliose occurring 
in the Amboina group, with the single addition of the mono- 
typic j\lel('»^icteris; but all the species, except one, are different. 
No genus is strictly confined to New Guinea, the nearest a[i])roach 
being Jleloin/cteris, wliich is known onl}- from t!.is island and the 
])ismarck Archipelago. No zoogeograidiical jirovince has so rich a 
Nyclimcne fauna, all four natural groups of this genus being repre- 
sented, and two of the groups (the cijdotis and aello groups) found 
nowhere else. Six species are autochthonous, viz. Pteropnx cpvlarivs 
(closely allied to Ft.macroiis from the .\ru Islands), Dobsonia minor 
(no close relative elsewhere), D. ma<jna (scarcely more than sub- 
specifically distinct from D. molvccensis from the Amlioina groiij) 
and Aru Islands), Nyctimene cyclctis and ccrtans (two closely allied 
peculiar species), and N. acllo (pec\iliar species) ; to these may be 
added Fteropus papunnvs (doubtfully distinct from Ft. nevhihernicvs 
from tlie liismarck Archipelago), and Ft. hypomdanus Intciis and 
jS'i/climene ffeniinvs, both of wliich have spread only to the small 
islands at the eastern eytremity of New Guinea ; making, at the 
vcrv highest, nine (piite or nearly autochthonous forms out of a 
total of sixteen. The dislribution of the seven forms not entirely 


confined to New Guinea is : (1) occurring both west (Amboiua group) 
and east of New Guinea (Solomon Islands), Hoiisetius hrachi/otis : 
(2) common with the Gilolo group, Pteropus chrysauchen: 
(^3) common with the Key and Aru Islands, hut not extending to 
Australia, Macrofjlossus hujochilKS nanus: (4) common with Aus- 
tralia and also with either Key or Aru Islands (if not both), JSycti- 
mene papuanus and Suconiicteris crassa papuana : (5) direct invader 
from New Guinea through the Torres Straits islands into Australia, 
Pteropus conspicillaius : (6) common with the Bismarck Archipelago 
only, Melonycleris melanops. A faunistic contrast between the 
North-west and South-east of New Guinea is, so far, only indicated 
by Pieropus chri/saachtn of the former area (extending to the Gilolo 
group) being replaced in the latter by its geminate species. Pi. con- 
spiciUatits (extending to N. Australia) ; but more definite suggestions 
for a subdivision of New Guinea into faunistic districts may be 
derived from the distribution of the species of Nyctimene when 
better known. 

Key Islands. — Four genera have been recorded, Pteropus, Doh- 
sonia, Nyctimene, and Syconycteris, each (so far as known with 
certainty) represented by one form ; in addition to these, Macro- 
glossiis probably occurs in the islands. Of the four forms three are 
ncculiar, viz. Pteropus keyensis, a species allied to Pt. melano- 
pogoii from the Amboina group, Banda Islands, and Timor Laut, 
but with no close relative in New Guinea; Dohsonia viridis viridis, 
also with Amboiuan affinities (no ecjuivalent in New Guinea); and 
Syconycteris crcssa Jceyensis, which is more intimately connected 
with the New Guinean than with the Australian race of the species. 
The fourth form, JSyciiinene p)apuanus,\s common to the Key Islands 
and New Guinea. As evident from this, the Fruit-bat fauna shows 
both New Guinean and Amboinan affinities, and the former are in so 
far more intimate than the latter, as the two Amboinan forms have 
differentiated the one into a distinct species, the other into a sub- 
species, whereas of the two New Guinean forms the one is unchanged, 
the other distinguishable as a subspecies. 

Arw Islands. — Nyctimene has not been recorded from these 
islands, otherwise the genera are the same as in the Key Islands, 
but the relative distinctness of the Aru fauna is shown by the fact 
that no form is known with certainty to be common to both groups 
(Dohsonia inoluccensis and Macroylossus layochilus nanus may occur 
in both). The affinities point partly toward the Key Islands and the 
Amboina group (Pteropus aruensis, a peculiar species, related to 
Pt. keyensis and to the Amboinan Pt. melanopogon ; and Dohsonia 
violuccensis, indistinguishable from the Amboinan form), partly and 
more decidedlj- toward New Guinea (two common forms, Macro- 
ylossus lagochilus nanus and Syconycteris craesa papuana ; one 
distinct species, Pteropus macrotis, closely allied to the New 
Guinean Pt. ('pularius). 


Bismarck Arddpelago. — 7 genera, 8 species. Four (or, if 
Plerop us Jieohibc miens is distinguishable from Ft. papuanus, five) 
forms are peculiar. Naturally the fauna is chiefiy influenced by 
that of the neighbouring New Guinea ; the genera are the same, 
and of tlie species and subspecies liousetlus hracluiotis, Macroglossus 
Inyochilus nanv.s, Melomicteris melutiops., and (probably) I'leropns 
ntohihemicus are direct invaders, Niictiinene major and Syconycteris 
crassd finschi only slightly altered iudigenous representatives of 
JS'ew (iuinean forms. But at the same time there seems to be a 
distinct Moluccau (non-Papuan) element, represented by Pteropus 
cupistratus and JJobsonia pnedairi.v, both of which, though well- 
diHerentiated indigenous species, have their closest relatives in the 
Gilolo and Amboiua groups, but (so far as known) none in New 

Sohinon Islands. — 7 genera, 16 species. Whether the genera 
are compared with those of New Guinea or with those of the 
Bismarck Archipelago, the result is the same : Syconycieris is 
absent, Mdonyctcris is replaced by the closelj' related Nesonycteris, 
and Fteralopea- is added, otherwise there is no change. But the 
amount of autochtlionous forms is enormous, all beiug peculiar, except 
lionsettus hracliyoiis, and of the seven genera two (with altogether 
three species) are jjeculiar, viz. those just referred to, Fteralope.c 
and Nesonycteris. The sixteen species may be classed according 
to their probable faunistic and natural athnities as follows : — 
(1) Direct invader from west, Ilouseitns hracJn/olis, ranging from 
Amboina, through New Guinea and the Bismarck Archipelago, to 
these islands : (2) Indigenous representatives of New Guinean types, 
three, viz. Nyctimene scitalus, most closely allied to N.geminusivom. 
New Guinea and N, major from the Bismarck Archipelago ; Macro- 
f/losstis layochilus uiicrotus, the most closely allied races of which are 
found in tlie Bismarck Archipelago and New Guinea ; Nesonycteris 
u'oodfordi, as mentioned above the Solomon Islands representative 
of Melonycteris mr.lanops from New Guinea and the Bismarck Archi- 
pelago : (3) Indigenous species having their closest known relatives 
in the Moluccas only or both in the Moluccas and Bismarck Archi- 
pelago, but apparently none in New Guinea (compare remarks on the 
Moluccan element in the fauna of fclie Bismarck Archipelago) ; here 
belong, first, five species of Fteropus, viz. Ft. grandis, laveUanvs, 
rnbianus, rayneri, and coyuaius, all representatives of the Ft. ray- 
neri group, the only other known species of which inhabits the 
Amboina group ; second, the two species of Dohsonia, D. inermis 
and MfSOT, representing a group of the genus inhabiting the Amboina 
grou]), Gilolo group, and Bismarck Archipelago, but not known to 
occur on the continent of New Guinea : (4) Indigenous species 
having their closest relatives in Australia, three, viz. Fteropus 
tvoodfordi, a species strikingly similar in all essential characters to 
the peculiar Australian Ft. scapulalu.t, only much smaller; Fteropus 
culoHus and ^olomonis. two species of the Ft. hj/ponielanus group 
and both perhaps most closely related to the Australian Ft. brmineusi 


(5) Indigenous representatives of a chiefly Polynesian type, th'o 
peculiar genus FleraIo2'>ex (two species), allied to the I'teropiis 
psehiplion group, tlic members of which are North-west Polj-nesian 
in distribution, with the exception of one species occurring in the 

A discussion of the distribution of the various species throughout 
the Solomon Archipelago would to a great extent be prematv;re ; 
the Fruit-bat fauna of northern chain of islands (Ctioiseul, Isabel, 
Malayta) is as yet entirely unknown, that of the extreme eastern 
islands (San Christovnl group) very imperfectly explored, and there 
is probably not a little to be added to our knowledge of the Eruit- 
bats of the other islands as well. But two important facts are 
already now sufficiently well established to call for some comment 
here. First, that some species (liouseiiiis brachyotis, Niictimene 
scihdus, Nesonycttris woo'lfortli, and probably Pteropus ivoudforili) 
are evidently uniformly distributed from the Bougainville group in 
the west eastward through the southern chain at least as far as 
Guadalcauar. Second, that in spite of this spreading of a few 
species over the greater part (if not the whole) of the Archipelago, 
there is unmistakable evidence that the Fruit-bat fauna of the 
Solomon Islands consists in reality of a series of more or less dis- 
tinctly separated " fauuula\" This is most clearly shown by the 
distribution of the si)ecie8 of the Pteropus rayneri group : Pt. yrandis 
inhabits the Bougainville group, is in Vella Lavella replaced bj- 
Pt. lavellanuK, in the New Georgia group by Pt. ruliiamis, in the 
Guadalcanar group by Pt. rayneri, and in the San Christoval group 
by Pt. coynatns. Other, though much less complete, evidence to the 
same effect is afforded by the two species of the Pteropus Jiypo- 
meJanus group {Pt. colonus in the Bougainville group, replaced by 
Pt. sohmonis in tlie New Georgia groui>), the two species of Ptera- 
lope.v {anceps in the Bougainville group, replaced by atraia in the 
Guadalcanar group), and the two Dobsonia (nesea in the Bougain- 
ville and New Georgia groups, replaced by inermis in the San 
Christoval group) [compare the distribution of the three known 
Solomon Islands species of the Microchiropteran genus Hippiosiderus, 
IJ. diiwps in the New Georgia group, H. oceanitis in Guadalcanar, 
and //. deinissus in San Christoval]. The faunistic areas of the 
vSolomon Archipelago indicated by the distribution of the five 
species of the Pteropms raj/neri group are very nearly the same as 
those lately recognized by ornithologists. 

Australian subregion (Continent of Australia). 

o genera (none peculiar), 8 species (six peculiar). The only 
genera which have reached the Australian continent are Pteroptis 
(five sjiecies), Nyctimene (two), and Syconycteris (one); all of these 
occur also in New Guinea, which however is inhabited, in addition, 
by four genera which have not spread to Australia, viz. liottsettus, 
Pohsonia, MacroyJossus (extends to the Torres Straits islands, but 
has thus far not been recorded from Australia), and Mdonycteris. 


As might be expected New Guinean affinities are predominant, no 
less than five of the eight 6])ccies pointing in their origin, eitlier 
nndouhtcdiy or at leiist ])robably, toward the great neighbouring 
island ; PUrojms cons/ncillaivg and Nyctlmene pajmunKs are direct 
iinehan.i^ed inva lers from New Guinea ; Plerojnis polioccpludas is 
probably the Australian representative of Ft. ejinlarius (New 
Guinea) and Pt. macroiis (Aru Islands) ; Nyct'imene robhisonl, a 
ratlicr peculiar species, is perhaps most closely related to A', ganinus 
(New Guinea) and N. hdluJo' (Woodlark Island) ; and iiyconyderis 
auslralis differs only in -trivial characters from S. crassa, a species 
which, differentiated into several local forms, ranges over New 
Guinea and its satellites, west to the Amboina group. Two 
Australian species of Pkropus, scajndatus and hvunneus, have, so 
far as known, no equivalent in New Guinea, but related forms 
inhabit the Solomon Islands {luoodfonU; cohnus and nolomonis). 
And one species, Pteropus goiddi, points toward the Lesser Sunda 
Islands and Celebes, being very closely allied to Ft. cdecto. No 
Fruit -bat has been recorded from AVestern Australia, south of tlie 
Kimberley Division, or from Southern Australia, west of Melbourne ; 
and the single record (by Temminck, but contradicted by Gould) 
of a Fruit-bat from Tasmania {Pteropus ]JolioccpTudus) remains 
unsupported by recent evidence. 

Polynesian sulre<jion. 

Pteropus (18 species) and Notopteris (2) are the only genera 
represented. All species of Pteropus, as Avell as the genus JVoto- 
pteris, are pecirliar. The eighteen species of Pteropus represent 
live natural groiips of the genus : the Pt. luj pomeJanus group has 
s]iread only to New Caledonia (one species) and the Loyalty Ishinds 
(another); tlie closely related Pt. maviannus group covers (with 
seven species) ])ractically the whole of the subregion, so far as 
inhabited by Fruit-bats, except the lioniu Islands, and the group is 
entirely Polynesian, except for one species that has established itself 
in the South Liu-Kiu Islands; the Pt. Jombocensis group is represented 
only in the Carolines (one species) ; the Pt. pselaplton group ranges 
(with five species) over north-western Polynesia, from the Boniii 
Islands, south through the Pelew Islands and Carolines, to tho 
Santa Cruz Islands, and is again entirely Polynesian, e>icept for 
one species inhabiting the Pl\ili])pincs ; farther south and east in 
Polynesia (New Hebrides, Fijis, Samoas) it is replaced by the 
related and purely Polynesian Pt. samoensis group (three species). 
Each of the eighteen species is (so far as knowu) confined to one 
group of islands, except Pt. tjeddiei, which is common to the New 
Hebrides and the neighbouring New Caledonia, and Pt. ionymnis, 
which is common to the Fijis, Tongas, and Samoas. The autochtho- 
nous genus Notoj^teris has its closest relatives in the Solomon Islands 
(Nesonyctcris) and the Eismarck Archipelago and Ncm' Guinea 
( Melon ycteris) : it has thus far been recorded from the Carolines in 
the north, and New Caledonia, New Hebrides, and Fiji Islands in 
the south. 


Indications are not wanting that the Megachiropteran fauna of 
the Carolines, spread as it is over many islands covering a large 
area, may fall into at least two sections. Thus, the Pteropus 
marianyius group is in the extreme west represented by one species 
(yapensis), in the extreme east by a clearly distinct species 
(ualanns) ; iihe Pte7'opus j^selaphon gronp has so far been recorded 
only from two places in the Carolines, the Ituck Atoll and Mortlock, 
but specimens from the former {insularis) seem to differ from the 
single available specimen from the latter (phceocephalus). 

v. sruopsis of the sttbfamilies akd genera of 

I. Molariform teeth not multicuspidate ; lower 
canines not proclivous. 
A. Tongue simple : tixed to floor of mouth 
by posterior half, and without uufriuged 
filiform papillse at tip (figs. 65 A-H, [p. 1. 

p. 725) I. PTEROPODIN^, 

a. Facial axis of skull conspicuously de- 
flected against basicranial axis : alveolar 
line, if projected backward, passing 
through middle or upper edge of occipi- 
tal condyle or even through some point 
of supraoccipital [Rousetti;s section.] 

«'. Incisors never j — ^, lower canines nor- 
mal in position, cheek-teeth ^ or (in 
Styloctenium only) ^ ; cranial rostrum 
not shortened, premaxilla? not more 
reduced than usual ; second finger 
a'. Occipital (sublambdoid) region of skull 
not elongate ; a short tail (third meta- 
carpal nearly always slightly but dis- 
tinctly longer than fourth and fifth). 


«'. Incisors | — -. 

«*. Tympanic forming a short bony 
auditory meatus, premaxillsa 
spaced in front ; m, equal in 
length to mj and m, combined. 
[3 species: Ethiopian and Mala- 
gasy regions.] 1. Eidolon, p. '2. 

h*. No bony auditory meatus, pre- 
maxillse in contact or co-ossified 
in front; m, shorter than mj and 
mj combined. [14 species : Ethio- 
pian, E. Mediterranean, Oriental, 

Austro-Malayan.] 2. Rousettus, p 10. 

6\ Incisors -^ — ^ (i' lost), premaxillse 

spaced in front. "1 species: Celebes.] 3, Eoneia,^. bb. 


l'. Occipital (sul)laiubdoid) region of 
skull elongate, subtubular; no tail 
(tifth metacai-pal nearly always 
slijxlitly but distinctly longer than 
third and fourth. [Pteropus sub- 

c^. Incibors | — ^, cheek-teeth g. 
c\ rrenia.xilliB in simple contact in 
frout ; io once and a half to .six 
times the built of i,, upper canines 
without secondary cusp (e.\cept in 
I'feiopus tithe) culadts). 
a'. No well-ditt'ereutiated antero-in- 

ternal tubercle in p' and m', no 

sharply delined inner basal ledge 

in p^, ni|, and m^ (^except in Pt. 

imetianits). [85 species, 103 forms: 

Malagasy, Oi-iental, Austro-Ma- 

liiyan, Australian, Polynesian ; 

one Ethiopian species.] 4, Pteropus, p. 61. 

l>\ A well-differentiated antero-in- 

ternal tubercle in p* and m' (some- 
times also in p^ aiid -p^), a sharply 

defined inner basal ledge in p^, m„ 

and m„. [6 species, 9 forms : 

Philippines, W. Austro-Malaya.] o. Acerodun, p, 41:.'. 
dK Premaxilla? co-ossified in front (in 
adults) ; i, twelve to fifteen times 
the bulk of i,, upper canines with 
large external secondary cusp, 
upper niolariform teeth with promi- 
nent anterior and posterior basal 
ledges, external main cusp of p^, 
m,, and m^ distinctly bilobed. 
[2 species : Solomon Islands.] .... 6. Pferalope.v, p. 432. 

2 2 -,5 

(P. Incisors j — ^ (i, lost), cheek-teeth j 

(nij lost). [1 species: Celebes.] .. 7. Styloctenium, -p. AA2, 
U. Incisors j — j (i' and i, lost), lower ca- 
nines at extremity of mandible, cheek- 
teeth y (p^ lost); cranial rostrum at 
least slightly, often considerably 
shortened, premaxillae sublinear ; 
second finger without claw, a short 
tail, wings from spinal line, back 
seemingly naked (the furred back 
being covered by the naked wing- 
membranes). [12 species, 13 forms: 

Austro-Malayan.] 8. Bobsonia, p. 448. 

b. Facial axis very little deflected against 
basicranial axis : alveolar line, if pro- 
jected backward, passing through lower 
edge of occipital condyle or even some 
distance below condyle (exceptions : 
Plerotes and 82)h<trici»). 


/. Biaiii-case peculiarly flattened pn3tc- 
riorly ; a small whitish hair-tuft at 
base of ear-couch anteriorly and poste- 
riorly (except in Scotomjct.eris). [All 

genera Ethiopian.] [Epomophoeus 

C-. Cheek-teeth \ (ur absent), molariforai .siicxio.vf.] 

teeth sublineav, with flattened crowns ; 
palate-ridges simple (tig. 29 J3, p. 48-3). 
[1 .species.] P. TkrMs, p. 483. 

d-. Cheek-teeth g fp^ m", and m.^ aV- 
sent) ; some or all of the palate- 
ridges peculiarly modified, 
e^. Cranial rostrum long : orbit to tip of 
nasals much more than lachrymal 
e*. Rostrum long and broad, post- 
dental palate flattened posteriorly. 
<■'. Koslrum not deeper than usual, 
premaxilla; in simple contact 
antei-iorly ; lower incisors and 
canines biting against corre- 
sponding upp'^r teeth, outer ridge 
oV lower molars simple ; muzzle 
without cutaneous leaves, tail 
rudimenfary (two free caudal 
vertebrtie) ; males with shoulder 
pouches and erectile shoulder 

brushes. [3 species, 4 forms.] . . 10. Eiuy.nop:', p. 487. 
d"^. Rostrum considerably increased in 
depth, premaxiilje ankylosed to- 
gether anteriorly ; lower incisors 
and canines closing some distance 
in front of upper, outer ridge of 
lower molars bilobed or trilobed; 
upper lip with cutaneous leaves, 
tail absent (no free caudal ver- 

tebraj); no shoulder pouches or [p. 501. 

brushes in either sex. [1 species.] 11. Ihjpsiynathus, 
/•'. Rostrum long and narrow, post- 
dental palate deeply depressed [p. oil. 
posteriorly. [8 species, 9 forms.] 12. Epomophorm, 
f^. Cranial rostrum short: orbit to tip 
of nasals equal to or less than 
lachrymal breadth. 
^'. I'ostzygomatic palate not (or 
scarcely) narrower posteriorly than 
anteriorly ; bead %%-itliout white 
markings (except for the usu.'il 
Epomophorine whitish tufts at 
base of ear-conch anteriorly and 
* e\ Postzygomatic palate at least as 
long" as broad ; maxillary tooth- 
row extending backward very 
nearly to ventral margin of orbital 

synopsis OK fJENKRA. XCV 

cavity; soft palate wiih a deep 

median ^-oove, narrowiujj puste- [p- •^•jA. 

riorly. [1 species.] ,13. M'uropteropics, 

/■'. Postzygomatic palate almost twice 
as broad as loi;g ; maxillary tooth- 
row not nearly reacliiug ventral 
margin of orbital cavity ; soft 
palate with a prominent median 

keel. [1 species.] 14. Xaiio/ii/rt>ris, p. 559. 

/('. rostzygomatic palate either much 
uarrow-er posteriorly than ante- 
riorly or entirely absent ; head 
with poculifir white markings. 
[/''. Postzygomatic palate long, its 
lateral margins forming straight 
lines converging posteriorly; no 
whitish tufts at base of ear-conch. 

[1 specie^-.] 15. Scotoui/cferis, p. 563. 

k'. N postzygomatic palate, the meso- 
pterygoid fossa continued forward 
nearly to level of posterior molar ; 
small whitish hair-tufts at biise 
of ear-concli anteriorly and poste- 
riorly. [1 species.] . .' 16. Cti.uiii/clen's, p. 568. 

<J'. Jh-ain-case not flattened posteriorly ; 
no vvhitisli tufts at base of ear- 
conch [Cynopteeus sp:ctiox.] 

e'. Cheek-teeth j;; rostrum less shortened: 
orbit to tip of nasals subeq-ial to 
lachrymal breadth. [The only Ethio- 
pian genus of the section : 4 species.] 17. Mtjomjcteris,^.b7Q. 

f-. Cheek-teeth less than ^ ; rostrum 
much shortened ; orbit to tip of 
nasals much less than (only in 
Sphcerias subequal to) lachrymal 
(f. Two or one pair of lower incisors, 
lower canines normal in position ; 
no3trils not tubular ; tongue witli 
three circumvalhite papillse. 
»'. Posuirbital foramen (through base 
of pojtorbital process) present. 
!\ Postorbital foramen large, pre- 
maxilbe in simple contact in front ; 

cheek-teeth j (m- and m, lost) ; 
membranes from first toe. 
«''. Cranial ro.^trum much lower at 

canine than at p^ ; upper canines 

^vith .'^econdajy cusp at inner 

edge ; tail present. 

o". Incisors 7, — ^, i" not shortened ; 
upper canines without vertical 
groove on front face. [6 species, 
1(5 forms : Oriental, \N . Austro- 
Malayan.] 18. Cynopterus, p. 08G. 



L\ Incisors j— j (ii lost), i' much 
shortened ; upper canines with 
distinct vertical groove on an- 
tevo-medial siu-tace. [I species: 

rhilippines.] 19- Ptenchirus, p. 643. 

t". Cranial rostrum as deep in front 
(at canine) as at p'' ; no secondary 
cusp in upper canines; tail ab- 

sent. [1 species :Indo-Malaya.] 20. Mer/^rojjs, \x bib. 
j\ Tostoibital foramen extremely 
small ; prenia.\^illae ankyloeed to- 
gether in front; cheek-teeth = 
(also p' lost); membranes from 
second toe, tail present. [1 

species : Indo-Malaya.] 21. Dyacoptems, p. 6ol. 

J*. Postorbital foramen absent. 
k". Incisors simple (terete, crown not 
peculiarly differentiated), sub- 
vertical, cheek-teeth not narrow ; 
interfemoral and calcar unmodi- 
c". p^ with autero-exterual basal lube 
or cusp, 
c" Pren)axilhe in simple contact 

2 2 

anteriorly ; incisors ^— •[ (i, 
lost), i' much shortened, cheek- 
teeth \ (only m3 lost), p' with 
lar^e antero - external basal 
. , lobe; wings spotted. C^ ^ _ ,. , . (.-, 

I ■■'■■■ ' species: Borneo.] 22, Bahonycten., p. 6-4. 

(V. Premaxillre solidly united au- 

2 2., 1 

teriorly; incisors^— 2,iiiorm at, 

cheek-teeth \ (m^ and m'* lost), 

p^ with well-defiued antero- : 

external basal cusp; wings 

(probably) not spotted. [I ^ ^, . p^^ 

species : Java.] . ^3. Clwonax, p. 0o8. 

d". p' unmodified. 

e'. Incisors |— g, i'^ normal, p^ and 

nil "^ith surface cusp: tail 

rudimentai-y,wing3 from second 

toe. [1 species: W. Austro- 

Malaya.] 24. Thooptems, p. 662. 

f\ Incisors \—\ {h lost), i* much 
sbortened, no surface cusp in 
any cheek-tooth ; tail about 
half as long as foot with claws, 
wings from first toe. [1 species: 

Indo-Malaya.] ._^. • • • 25. renthetor, p. 665. 

l\ Upper and lower incisors (^— ^) 

proclivous, croM-ns highly differ- . ' . 

entiated, triangularly pointed^ ^^ , ^^^^ 


clioiilc-ttPtli (-) iiniisiinlly narrow; 
iiit<'rrt'iiioral imt fxteiulinr!- be- 
vdiul miildlc cil' tibia, calcar and 
tail al)sent. [1 .«pecies : Inclo- 

A^ Incisors „— ,, (i-, i,, i., lost), clieelc- 
tf^eth j (ni-, m., lost), lower canines 

close tog-etlier at extremity of 
mandibh ; nostrils elongated into 
cylindrical tubes, tcnignu with four 
circuin vallate pa]iill!c, tail not much 
shorter than tibia, winjis spottei]. 
[13 spcjies : Anstro-Malaya, Ans- 


I. Tonn-ne more extensible, fixed to floor of 
mouth by its postei ior third, its terminal 
fourth or fiftli coveied above with iin- 
f'rinuvd hlifornr papilhie (figs. 60 I-L, 


f. Premaxillm not broader in upper than in 
lower half', infraorbital canal slun'ter; 
third metacarpal longest, or third. 
f'liuilh. and ti'th siibequal, terminal 
plialanx of third fing'er always mnr-li 
sjiorter than tliird metacarj)al. fp' 
and )), often unnsnally larj;e, nevtr 


e'. I'remaxilhe in simple eonta<t in front 
or slightly spaced ; cheek-teeth less 
reduced in size : second finger without 
claw, tail snbequal in length to foot 
with claws. (Facial axis not strongly 

deflected ; incisors f, — t, cheelt-teefh L- 
or, by suppression of small m,, ",.) 
[:i species : Indn-]Malaya, Celebes.] . . 
/'. I'reniaNilhe solidly fused in front ; 
clie(>k-teetli much more reduced ; 
second finger clawed, t.iil rudimentarv 
or absent. 
//-. rrpper incisors minute, crowns not 
peculiarly diHevenliatfcd, lower in- 
cisors subequal in size ; interfemoral 

and calcar unmodified. (IncisorsF, — ", 
cheek-teeth '^, but individual anoma- 
lies in number of cheek-teeth of 
fVeipient occurrence in Macrnt/losK>i.<.) 
P. ]'' axis much lesj deflected, pre- 
maxilho less pi-oclivous ; p' (second 
premolar) much higher than ]>', 
l()wer incisors bilobate, subeqnaliy 
spaced ; fifth metacarpal nuu-li 
shorter than third, fourth inri'r- 
mediate; males ivith 1-irge neck- 
tufts. [1 species: I'Uhlopiuu.] . . . . 

2(j. S/i/ur-r/ds, p. G71. 

Nijcfimnif, p. 6yl. 

[SIN/K, p. 723. 

oxvrTiaus siu'Tion. 

'2i>. JSo7n;c'i'i/\. p. 7i.'8. 

29. Me.^a'o(ilossus, 

[p. 738. 


/'. Facliil axis stroiifrly deflecled, pre- 
maxillre more proclivous ; p' more 
reduced, little hijjbev than pS 
lower incisors simple, i, — i^ widely 
spaced; third, fourth, and tiftli 
n?etacnrpals subequal ; males with- 
out neck-tufts. | "2 species, 6 forms : 

Indo-Cliina to Solomon fslandf.] . . 30. Macroyhssm, p. 746. 
h-, UjSper incisors large, crown well 
dilierentiated, narrowly chisel - 
shaped, i,, mucli larsjer than i, and 
with obliquely triangular crown ; 
lateral interfemoral unusually nar- 
row, calcar rudimentary. (Incisors 
^ — 5, cheek-teeth '^ or, by suppression 
of small nij and ni^, ..) [3 species, 

7 forms : Austro-Malaya, A.ustralia.1 31. iSi/co77i/c/cn's, \^. 771. 
(J. PremaxilliB twice or thrice as broad in 
upper as in lower half, infrairbital 
canal long-; fifth metacarpal lonirest, 
terminal phalanx of third finger sub- 
equal to or longer than third meta- 
carpal, (p' and p, minute or absent.) . [Notoptf.kis skction.] 
[/' . Premaxillre in simple contact in front 
or slightly spaced, anirular process of 
mandible large ; p' and pi present 

(cheek-teeth '^), p, smaller than m^ 
and m^ : tail absent, membranes from 
flanks and third or fourth toe, back 
furred, tibia much less than half the 
length of forearnj. 
{-. Incisors | — 7,; second finger clawed. 

(Underparts much darker than lack.) 

[1 species: New Guinea, liisuiarck 

Archipelago.] 32. Melonyderis 

j-. Incisors y — ^ ('i ^o^^) ' f^econd finger 
without claw. (Underparts paler 
than back.) [1 species: Solomon 

Islands. | 

//. Premaxillae co- ossified in front, an- 
gular process of mandible small ; p' 

and pi absent v'cheek-teeth -^), incisors 
- — " (or in adults, i' being deciduous, 

— ), p^ the largest lower cheek- 
tooth ; tail very long, snbeqnal to 
forearm, membranes from spinal line 
and first and second toe, back seem- 
ingly naked at middle (the furred 
back being covered by the naked 
membranes), second finger without 
claw, tibia longer than usual (half as • 
long as forearm). [2 species : Poly- 
nesia.] ■ 34. Nofopferh. p. 793. 

p. 78.') 

33. Kesoin/cferis, p. 790. 


II. Molarifonii teeth inulticus|iidfite, loTrer 
canines strongly proclivous (crossing- 

upper canines at nearly rifjht angles) III. IIARPYIO- 

I'remaxillaj!ietl in front ; incisors j — ; IL il'^lilA-r., p. / .*• . 
(i' and i, lost), lov.'ev pair rudiuseu- 
tary, uppei' canines bieuspidatej lower 
canines tricuspidate and situ-ited close 
together at extremity of nuibdible, cheek- 
teeth ^ ; second finger clawed, tail absent, 

tibia uuusiially short. [1 species: Philip- [p. 7i).). 

pines.] , .'{5. Harpi/imii/rferis *, 

" Knj/ " to the (jenera, husrd cniirelij on dcntid atul cmnidl 

Ill the subjoined artificial "key" to the genera such characters of the skull ami 
dentition have been selected as are easily observable auil readily understood b\- 
anybody witb an elementary knowledge of Maunuuiian osteology and odontoloy,y. 
liut the siniplitication of the key bas liad to be l)Ouabt by sacrifice of the seriid 
ariangenient of the genera adopted in this Catalogue. [A complete key to the 
genera of Megachiroptera based entirely or chiefly upon their external characters is 
an impossibility ; a large number of genera, even though they may belong to widely 
distinct groups of tlie (Suborder, cannot lie discriininatud with certainty from eacli 
other without an examination of tlie skull uiul dentition.] 

a. No lower incisors JS'jc/imeiu; p. 081. 

ii. One or two pairs of lower incisors. 
a'. One pair of lower incisors. 
«-'. One pair of upper incisors. 

<(■'. Cheek-teeth ".on each side llitrpi/iuiii/cfcris, p. 799, 

h^. Cheek-teeth i on each side /J.///,soj/./(/, ii. 118. 

' * 

t-. Two pairs of upper incisors (middle pjiir de- 
ciduous in Notopteris). 

c'K Cheek-teeth '. on each side iWsoHmVici.s-, p. 790. 

(/•'. Cheek-teeth less than j, on earb side. 

<■(•' Cheek-teeth .. 

«\ Length of rostrum (orbit to tip of im.sals) 

greater than lachrymal breadth (across 

lower edges of hicbrymal foramina); 

111' (penultimate upper cheek-tooth) equal 

in size to yi^ (iintepenultiinate) UtijloctcuiKiii, p. 112. 

b''. Length of rosliiim much less than 

Ih( lirymal breadth : ni' much smaller 

than p' Balitiii/iftcrit:, \'. iibi. 

b'. Cheek-teeth t. 

(■■'. Hostruni long, strongly deflected: lowir 

incisors widely spaced ; reduced luimber 

of cheek-teeth due to loss of anterior 

premolar above and below (p' and p,) . . Xolapteris, p. 793. 
'!■''. Ivostrum short, scarcely deliected ; lower 

incisors close together; ledured number 

of cheek-teeth due to loss of posterior 

molar above and below (m- and iii^). 
<(''. I'ostorbita' foramen (through base of 
postorbital process) large. 

* Ou the affinities of IIar/)i/iniii,rttr>f. sec p. SUiJ. 


a'. SurCace cusps in in and ui] (third 
and fourth lower cheek-tooth; ; upjier 

canines grooved I'tciK/vhlrus, p. 613. 

b^. Ko surlaco cusps in check-'tteth ; 

upper canines without groove Meyarops, p. 154(3. 

t". Post orbital foraiuen absent J:'entl(etvr, i>. Wo, 

b'. Two pairs of lower incisors. 

C-. ihm pair of upper incisors Boneia, p. 55. 

rf-. Two pairs of upper incisors (outer pair often 
deciduous in Mpuniops). 
£••*. o upper cheek-teeth on each side (anterior 
tootii otteu deciduous in i'tempus and 

c^. A short bony auditory meatus Eiilvlvii, p. -. 

d''. No bony auditory uieatus. 
e^. Occiput elongate, subtubular. 

c^. Outer lower incisor not more (but 

g."nprally less) than six times the 

bulk of inner. 

c'. No well-defined an tero - internal 

tubercle in p'' and in' (third and 

fourth upper cheek'tootli) Fttni^nis,}'. t'A, 

d'. A well -defined antero - internal 

tubercle in p^- and m' Arcrodmi, y, 112. 

d''. Outer lower incisor twelve to fifteen 

times the bulk of inner I'fcralDpe.v., \u Sni'2, 

f''. Occiput not elongate. 

«'". Length of rostrum (orbit to tip yf 
nasals) subequal to lachrymal breadth 
(across lower edges of lachrymal 

foramina) M,ijouijCtel-is, \^. o~G, 

f'\ Length of rostrum conspicuously greattr 
than lachrymal breadth. 
ۥ. Outer lower incisor much higher than ; p. 771. 

inner .Svco/(//;-<c'c/s Ipt.i, 

/"'". Lower incisors subeciual in height. , ,, ,, .> ,,. 

rt", Clieek-teetli not linear •< -,, , -^ -.,^, 

6». Cheek-teeth linear. ^ di..M,jjru ■,■!>> ■ , p. /.s. 

«". Premaxilla' much broader in 

upper than in lower half IiUhnij/vtcris, p. 7tio. 

6''. Premaxilhe not broader above 
than below, 
a'". Lower incisors subeiiually 

spaced ?Jt;/'iln'//iiti!ii(s, p. 738. 

b'". Inner pair of lower iiici?ors 

widely spaced Maci-Kiilosiua. p. 7J<'. 

,/"''. \jes> than o upper cheek-teeth on each side. 
(,-■'. 4 upper cheek-teeth on each side. 

ij". t) lower cheek-teeth on each side I'Urtitci-, ji. 483. 

h^. o lower cheek-teeth on each side. 

/)''. Postorbital foramen (through base of 
postorbital process) large. 
J'. Rostrum short, scarcely deflected ; 
inner secondary cusp in u)iper 

canines ('j/uujjffru.':. \i, oS<^. 

h'. liostrum long, strongly deflected ; no 

secondary cusp in canines Hijconi/ctci-is (jit.). p. 771. 

Ifi. Postorbital foramen absent. 

/''. Incisors proclivous, highly differ- 
entiated, triangularly pointed Sphferias, p. 671. 

* As pointed out elsewhere (p. 733), it would be difficult to give any reliable (and 
practically useful) cranial or dental character by w hich Euui/ctefis spelcea and iiicijnr 
(K. ronfiiherrfi differs by the loss of m:i) can be discriminated from cnij/ s])ecies uf 
lioHsetlns. although these two genera belong tc difleicnt .subhimilies. Externally 
they are easily distinguished {Suusetlus with. Eonj/clcris without, claw in second 


J', Incisors subverticiil, simple. 

k'*. Surface cusp in third ami fouitli 

lower clieek-tooth (p4 ami iiiv) . T!ioiiplerit.t, \> 002. 
(l\ No surface cusp iu any clieek- 

tooth Cliirututj:, p. (j58. 

f^s 3 upper [and 5 lower! clieek-teeth on each 
(■•'i Length of rostrum (orbit to tip of nasals) 
much griater than lachrymal breadth 
(across lower edges of lachrymal 
t''. Post dental palate deeply depressed pos- 
teriorly Epiiimiji/iui-iWi \K oil: 

/•>. Postdental palate flattened posteriorly. 

X''. Outer ridj^e of lower molars simple ... Eputimps, \t. IS7. 
IJ. Outer ridge of lower molars bilobtd 

or trilobed UjjpsiyiwthaSi \). 'y)l. 

J'', Length of rostrum less than or subequal to 
lachrymal bre.idtli, 
jI^ Palate not extending beyond tootli- 

rows ^choanie at level of la^t molar) ... Cuaiiij/'.-lci'i.s, p. 5(i8. 
/'', Palate extending behind tootli-ro«s. 

m''. Premaxilla; fused anteriorly Dj/acvjitenis, \i. Hoi. 

ii/. l'r( inaxilla; separated by suture ante- 
c'*. I'ostzygoniatic palate much nar- 
rower [losteriorly than anteriorly . Sr(itiiii,i/(ieris, p. o''3. 
y*. Post/.,\gon!atic jialate subequal in 
breadtli throughout. 
c^. Upper tooth - rows extending 
l)ractically to ventral margin 

of orbital cavity MirrojiternpiiSy p. ooi. 

d'^. Upper toolh-rows not nearly 
reaching ventral margin of 
orbital cavity l^'unuiiyclcvie, p. .1.59. 


Differential characters, as compared with tlie suborder Micro- 
oliiroptera. — Second digit: retaining an evident degree of inde- 
pendence from third, its ungual phalanx present, though the claw 
is sometimes wanting. Deltoid crest of humerus low ; tuhercidani 
mnjits and t. minus small, the former never articulating with the 
scapula. Facial portion of skull vnriable in length, hut never 
specialized in form ; cochlea small, basioccipital and basisplieuoid 
not narrower than usual ; glenoid fossa of sqiramosal unmodified ; 
angular process of mandible broad and low, or practically absent. 
External ear not specialized, tragus never present, margin of ear 
forming a complete ring. 

Range. — Tropical and subtropical portions of Old "World, includ- 
ing the whole of the Malagasy region, and extending to Australia 
and Polynesia, as far east as the Carolines and Samoa, but not to 
Xew Zealand. 

Fnniily PTEROPODID.i:. 

Sul)faiMily I. PTEROPODIN.E. 

Differi'ittiiil cliaractfrs. Tongue not highly extensible. liioi.-o;j 

Ramie. — The same as tliat of the Megachiroptcru [lupra). 

1. EIDOLON, liojin. 

Ci/non)/cicris (pt.), Dobson, Cat. Chir. B. M. p. 70. 

1815. Eidolon, Rafinesque, Anah/se de la Nature, p. 54 . . E. lielvum. 
1861. Pterocyon, Peters, MB. Akad. Berlin, p. 423 E. helvum. 

1881. Leipouyx, Jentink, Notes Leyden Mies. iii. p. 60 (Jan. 

1881) \_nec Liponyx, Vieillot,18\Q, a genus of birds]. E. helvum. 

1882. Liponyx, Forbes, Zool. Record, xviii. (lor 1881), 

Mamin. p. 13 (nom. emend.). 

Diaf/nosis.- — Basicranial axis di.stinctlv deflected ; occiput not 
elongated and tubular ; a short bony auditory meatus ; palate 
much broader posteriorly than between canines ; length of rostrum 
much greater than lachrymal width ; front of orbit vertically 
above middle or posterior halt of m' ; premaxillaries separated in 
front. Incisors^ — ^; cheek-teeth ^ ; p' in cross section much 
larger than an upper incisor ; m^ equal in length to m, and m^ 
combined. Second digit clawed ; membranes from sides of back ; 
a short tail. Forearm 114-132 mm. 

Skull (fig. 1).— Brain-case moderately deflected : alveolar line pro- 
jected backward passing very nearly through bases of posttympanic 
and paroccipital processes and upper margin of occipital condyle. 
Occiput not (as in Pierojms) produced backward and downward 
into a conspicuous tube. Tympanic elongated externally into a 
short tubular bony auditory meatus (a peculiarity unique among 
Bats). Palate broadest between m'^-m^, breadth at palation border 
much greater than between canines, and about equal to breadth 
between inner sides of p* (cf. Pteropvs). Rostrum long : length 
from front of orbit to tip of nasals much greater than lachrymal 
■width of skull, subequal to (a little longer or shorter than) length 
of maxillary tooth-row ; front margin of orbit vertically above 
middle or posterior half of m^ (cf. Myonycteris). Tip of nasals 
nearly vertically above front of premaxillaries (cf. Pteropiis). 
Premaxillaries slightly, but distincti) and constantly, separated in 
front (cf. Boiisfttus, Myonycteris, Pteropus). Width of frontals at 
interorbital constriction slightly less than (or at most equal to) 
width at postorbital constriction. Postorbital processes in aged 
specimens reaching about halfway between frontal and zygoma. 
Sagittal crest short and low, the temporal ridges sometimes 
remaining .separate throughout the life of the individual ; larabdoid 
crest strong, projecting backward considerably beyond the plane of 
the supraoccipital. Posttympanic processes longer than paroccipital 
processes. Ectopterygoid processes small. 

i* i" c p^ p'' p* m^ m' 

Dentition (fig. 1). — r-. ^-^--^ X 2 = 34. p' and m% p, 

i,i^cp,p3p,m^m^_m3 ^ 'i' 

and m^ reduced. One skull of E. helvum in the collection (ad., teeth 

slightly worn ; no history ; specimen c' in the list — infrc'c p. 15) 

has a minute m"' on one side. 


Teeth without special modifications; no secondary cusps in 
canines or cheek-teeth. General structure of molars: a median 
longitudinal groove, flanked by a higher outer and lower inner 
ridge each ndge rising, or tending to rise, into a cusp in front of 
middle ot tooth ; in the upper molars the outer ridge representa 

Fig. I.— Eidolon helium, cT. Bilelipi, Fernando Po. No. \. 

cusps 4 and 5 in the typical molar in Insectivorous Bats, the inner 
ridge cusps 6 and 7 ; in the lower molavs, the inner ridge cusps 
1-3, the outer ridge cusps 4 and 5. 

Upper incisors small, terete, very nearly equal in size (the outer 
incisors, if anything, faintly smaller in cross section) ; crown 
indistinctly differentiated from shaft, subcircular in cross section, 

E 2 


cuttino;-e(lge bluut ; i'-i' widely separated (chiefly owing to 
separation of premaxillaries), the interspace being nearly equal 
to diastema i'^-c ; i'-i^ rather closely approximated. Canines 
simple ; cingjlum not sharply pronounced, no secondary cusps ; 
surface of crown as a rule marked by three shallow vertical 
grooves, one anterior, one internal, and one posterior. All post- 
canine teeth more or less separated; interspace c-p' much greater 
than diastema i"-c. p' small, but much less reduced than in 
Rousetlus, being in cross section at base of crown from three to 
six times the size of an upper incisor, and nearly equal to m^ ; as a 
rule situated nearer to the canine than to \)^ : as clearly seen on 
comparison with p' and p'', tlie crown of yj' is formed by the com- 
pletely fused cusps 4 and 6, a slight f sometimes obsolete) depression 
on its posterior face representing the remnants of the median 
groove, p' almost caniniform, its princi])al cusp (formed by the 
anteriorly completely fused cusps 4 and 6) from one half to two 
thirds the height of the canine, acutely triangular, sharply pointed ; 
a vertical furrow on its posterior face corresponding to the longi- 
tudinal groove in the posterior cheek-teeth, p"" molariform, longer 
(antero-posteriorly) than broad ; outer ridge raised into an olitusely 
triangular cusp, much lower than p"'' ; inner ridge forming a low 
cusp opposite the outer cusp; the inner bases of the cusps so 
closely approximated as to form a narrow, more or less complete 
bridge across the median groove, m' similar in structure to p^, 
but outer and inner ridge lower, more flatly rounded ; no indica- 
tion of a transverse bridge, m^ small, subcircular in outline, equal 
to about one sixth of m\ 

Lower incisors similar in form and size to upper ones, in contact 
with each other and with canines, or separated by minute spaces ; 
row slightly convex ; cutting-edges blunt, without any trace 
of a median emarsination. All postcanine teeth more or less 
separated. Pj small, from twice to four times the bulk of a lower 
incisor, but not reaching above level of cutting-edges of incisors ; 
as a rule situated nearer to the canine than to p^. Principal cusp 
of Pg (formed by the anteriorly completely fused cusps 2 and 4) 
about two thirds the height of the canine, acutely triangular, 
shar])ly pointed : posterior face of crown marked by a vertical 
groove (the remnant of the original longitudinal groove), p^ rather 
longer than broad; outer ridge raised into an obtusely triangular 
cusp, lower than p^ ; the opposite cusp on the inner ridge much 
lower, bluntly rounded ; anteriorly the outer and inner ridges are 
closely approximated, in some individuals completely fused (as 
in p ). m^ long, with rare individual exceptions equalling or 
exceeding the combined length of m., and m^ ; ridges lower than in 
y> , more flatl" convex ; sometimes a slight indication of a transverse 
liridge from the inner base of the outer cusp, m^ similar in 
structui-e to m,, but less than half the size of this tooth, m^ very 
small, subcircular in outline, equal to (or smaller than) Pj. 

PaJateridfjes. — In E.helvvm: 4-f3-H3, i.e. four anterior, un- 
divided, three middle, separated in the median line, three posterior. 

First ridge teiminatiug at oi- closely behind tlie canine ; second at 
or closely behind p' ; third at ]/ ; fourth at front of p* ; fifth at 
front of m' ; sixth closely behind ra" ; seventh far behind m^ ; 
eighth to tenth situated near jjalation border. Palate-ridges in 
E. sahceum and E. dupreanum not examined. 

Head. — Muzzle long, from front of eye to tip of nostrils almost 
etjual to distance from front of eye to base of ears. Nostrils sepa- 
rated by a deep groove ; inner margins moderately projecting. 
Inner margins of lips fringed with short pajnlliie. Median portioa 
of tongue with oblique rows of rounded papilla?, each papilla with 
three small, straight, 8harj)ly pointed, backwardly directed horny 

Ears. — Tij) of eai'-conch reaching hinder corner of eye ; outer 
margin miich more convex than inner, tip broadly rounded off. 
Autitragal lobe practically wanting. 

Wings. — Chief characters : second digit clawed ; wings from 
sides of dorsum, rather closer to spine than to Hanks, and from 
back of first toe. 

First phalanx of first digit twice the length of metacarpal. 
Second digit subequal to third metacarpal. Third metacarpal 
between six and seven tenths the length of the forearm, a little 
longer tlian the fourth metacarpal, which is a little longer than the 
fifth. First phalanx of third digit equal to two thirds of meta- 
carpal ; second phalanx subequal in length to metacarpal. First 
phalanx of fourth digit rather more than half of metacarpal ; 
second phalanx always somewhat longer than first. First phalanx 
of fifth digit less than half of metacarpal ; second phalanx subequal 
in length to first. See wing-indices, infra. 

First digit included in the membrane by the metacarpal and base 
of first phalanx. Notopatagium acutely triangular. 16-20 long 
fasciae in the plagiopatagium, viz. 4-G postanconeal, 12-16 pre- 


2nd digit. 

3rd digit. 

4th digit. 

5th digit. ! 



1 473 

1 ! 

1st 2-3 
ph. ph. 

141 128 
131 111 











668 369 

646 347 







Ist |2Dd 
ph. 1 ph. 

299 289 

272 284 


E. dupreanum 

E. hehumSi, tabaum. 



Tall. — Less than half the length of the hind foot ; basal portion 
connected with intcrfenioral by its dorsal integument, terminal two 
or two and a half vertebra; freely projecting. 

l>e.vual differentiation. — Adult males of all species have the 
glandular hairs on the foreneck and sides of the neck brighter 
coloured tlian females ; females of E. hdvuni and E. sabermn are 
lighter coloured above and below than males ; females of E. helvwu 
and E. sahceum (and dupreanum ?) average a trifle larger than males. 

Kanije. — Madagascar ; AtVicau coiitiuojit, from Stiiiiaar and 
Seiiegatnbia in the nort)i, to Xyasaland and Namaqiialand in the 
sonth ; S. Arabia. 

Halrhs.—On the Middle and r})per Nile, v. Heuglin found 
"■ Fleropus jjrt?»jan«H" (i.e. Eidolon helvuni) mostly in flocks, 
frequenting Bornssiis (ethioijicus. the fruits of which, together with 
those of Ficns and Curdia, seem to be their favourite food ; they 
sometimes literally eat themselves into the Borassus fruit, a shot 
bringing the fruit together with the bat to the ground. They fly 
by night as well as by day, their sight being apparently unimpaired 
even in the brightest sunshine, though their flight by day is some- 
what flickering and unsteady ; by night it is owl-like, straight, and 
they occasionally pursue each other, making rapid turnings with 
audible flapping of the wings. They are noisy and quarrel- 
some, alighting with great uproar on their roosting-places. — In 
Fernando Fo F. Newton found them feeding on the fruits of 
C'lrrica ixtpaya and Fersira gratlssima. — The British Museum 
specimens obtained by A. Whyte at Mt. Malosa, Nyasaland, were 
found suspended fiom the upper branches of tall trees, but " they 
also frequent caves in the rocks, v\ here they probably breed " ; those 
collected by E. Seimund in Fernando Po were mostly shot in palm 
or pLiutain trees ; ripe embryos or new-born young were obtained 
in Fernando Po by the same collector between the middle of 
February and the middle of March. 

E. sabceum was seen in considerable numbers near Lahej, Aden, 
early in March (Col. Yerbnry) and in the middle of August (A. B. 
Percival) ; they frequent the tops of the tallest trees, where they 
collect in large bail-like clusters of 10 to 50, but are by no means 
easy to detect ; " were it not for the characteristic chattering that 
they keep up incessantly, they would probably be overlooked 
altogether" (Yerbury). Percival found them very noisy in the 
roosting-places, "squeaking and swearing, making a great fuss 
eafly in the morning " ; they were apparently feeding on dates 
is hich (in August) were just ripe ; when the dates are ripening 
every bunch is put into a bag made of palm-leaves, for protection 
against these bats and the crows. 

Affinities. — Eidolon is related, though not very closely, to 
JloKsetius, with which it was united by Dobson in the genus 
•' Cynonycteria." It differs from Roiisetlns chiefly in the deve- 
lopment of a short bony auditory meatus, in the distinctly 
Repavated premaxillaries, the proportionally longer rostrum, the 
less reduced size of p', the unusually long m^, the smaller m^, the 
larger number of posterior palate-ridges, the increased number of 
fascife in the lateral wing-membrane, the much larger size, and in 
being restricted to the Malagasy and Ethiopian regions (in- 
cluding S. Arabia). In the larger size of p' and proportionally 
longer rostrum, it would seem to be more primitive than Rousettns, 
but it is on a higher level of development in the peculiar tubular 
lengthening of Ihe tympanic, the sejjaraliou of the preraaxillaries, 


the lengthening' of m^, the reduction of m^, the considerably larger 
dimensions, and the more pronounced sexual differentiation m 

The three species are closely interrelated. E. dupreanum, from 
Madagascar, witli its relatively longer rostrum and less modified 
fur structure, is apparently the least specialized form. E. sabcpum, 
from S. \V. Arabia, is a small-skulled and broad-toothed represen- 
tative of the African E. /lefvum. 

Nomenclature.— In 1810 (Ann. Mus. d'Hist. Nat. xv. pp. 90-98), 
E. Geoffroy divided the genus Ptcrojms into three sections, viz., 
" Roussettes sans queue " (Pt. e(htlis, edwardsi, vulgaris, ruhricoUis, 
griseus), " Iloussettes a queue " (Pt. stramineas, cfgyptiacus, amplexi- 
caudalus, marginatus, minimus!), and " Roussettes a ailes sur le dos " 
(Pt. paJiatus). Five years later (1815, l.s.c), Kafinesque raised 
these sections to the rank oP distinct genera, restricting the genus 
Pteropns to the species of Geoffrey's first section, i)roposing the name 
Eidolon for the second, and Pteronotus for the third section. 
Pteropus stramineus, Geoff, (i. e. VespertiJio vampyras helvus, Kerr), 
as being the earliest known of the tive species included by Geoffroj- 
in the section " Roussettes a queue," may be fixed as the type 
species of Eidolon, Raf. (Of. K. Andersen, Ann. & Mag. N. R. (8) 
i. p. 432 ; 1 May, 1908.) 

Sgnopsin of the Species- 

a. Rostrum relatively longer : front of orbit 

to tip of nasals equalling or exceeding 
maxillary tooth-row; fur longer, moii 
■woolly, not closely adpressed ; colour 
darker ; forearm 1'27-131 mm. (Mada- 
gascar) \. E. dupreanum, p. 7. 

b. Rostrum relatively shorter : front of orbit 

to tip of nasals less than maxillary tooth- 
row ; fur very short, closely adpressed ; 
colour lighter. 
a'. Skull larger: total length .54:-o-62-2 mm. ; 

tooth-rows longer : c-m''' 21-23'8 ; molars 

narrower; forearm 117"5-132. (African 

continent) 2. is. /teiuiim, p. 8. 

b'. Skull smaller : total length 51'6-5o mm. ; 

tooth-rows shorter: c-m* 19 2-208; 

molars broader; forearm 114-127. (S.W. 

Arabia) S. E. sabf^um, p. 1 o. 

1. Eidolon dupreanum, Schl. tj- Full. 

CynomjcUris dapreana, Dobson, Cat. Chir. B. 31. p. '/8. 

Cvnonycteris stramineus [nee Geoff.), Peters, MB. Akad. Berlin, 

'186H, p. 88o (Nossi B6). 
Pteropus dupreanus, Schlegel Sf Pollen, P. Z. S. 1866, p. 410 (N.W. 
Madagascar) ; iid., in Pollen ^- v. Dam,Rech. I'aune Madagascar, 
ii. pp. 17, 172 (1868: X.W. Madagascar). 


L'ynouyctfris dupreauii *, Peters, MB. Akad. _flf/7f«, 18(37, p, 866 ; 

Jentmk, Cat. Oateol. Mnmm. p. 264 (1887) ; id., Cut. Syst. Mavitn. 

p. 152 (1888) (types : X.W. ^ladao-ascar) ; Trouessm-t, Cat. Mamm. 

1. p. 85 (1897)'; Seabra, Jorn. Sci. Math. Lisboa, (2) \. no. 19, 

p. 159 1 (1898). 
Cvnoptenis (Xanthirpvia) dupieana, TrouesMirt, Rev. Si- Mag. Zool. 

'(3) vi. p. 207 (1878). 
( 'ynonycteris straminea var. dupreana, Barthit, P. Z. S. 1879, p. 769 

(S.E. Mada.L'ascar). 
Xantharpyia dupreaua, Matschle, Megachiropicra, p. 63 (1899). 
llousettus dupreanus, Trouessart, Cut. Mamm., Suppl. p. 59 (1904). 
I'terocyon diipivauus, K. Andersen, Ami. ^ Mug. X. H. (7) xix. 

p. 504 (1907); Miller, Fam. ^- Gen. Bats, p. 56 (1907). 

Differs from E. helvum in the following particulars : — Cranial 
rostrum relatively longer and slenderer: measured from i'ront of 
orbit to tip of nasals equalling or slightly exceeding the length of 
the upper tooth-row from front of canine to back of m"; posterior 
premolars and molars slightly broader ; m' comparatively longer, 
being conspicuously longer than p^ ; p^ and m„ comparatively 
larger; fur on body longer, more woolly, and not closely adpressed ; 
colour of fur browner (details see below) ; length of forearm as ia 
large sjKicimens of E. helvum, but metacarpals and phalanges rather 
longer, as shown by the wing-indices, p. 5. 

Distribution of fur on ears, limbs, and membranes as in E. helvum. 

Colour. — General aspect : brownish above, tawny olive below ; a 
tawny half-collar in adult males. — Upperside of head, back, and 
ribiae brownish, approaching Prout's brown ; base of hairs tawny 
olive. Glandular hairs on foreneck tawny, paler on the sides of 
the neck ; absent in the only female examined. Breast and belly 
tawny olive on flanks, washed with dark hair-brown in the middle. 

Measurements. See table, p. 16. 

Range. Madagascar. 

Cotgpcs (two specimens) in the Leyden Museum. 

«. 5 ad. rk. ; sliiill. Madagascar. E. Bartlett [C.]. 

b. cJ ad. sk. ; skull. Viuauitelo, Betsileo, Hoyal Society 

Madaffasear; 27 May, [P.]. 



c. c? aJ. st. ; bknll. Itanibelo, Betsileo, Royal Society 1* 

Mndagascai- ■. H June, [l^-]' 



'2. Eidolon helvum, Kerr. 
Cynongcteris straminea, Dobson, Cat. Chir. B. M. p. 77. 

Cliaiive-Soui'is, Des Marchais, I'uyage e>i Gidnee, i. p. 81 (1730: off 
Guinea Coast). 
" I/est-er Tomate Bat, Pennant, Sgn. Quadr. p. 362, no. 274/3, pL xx.xi. 

* \ ariuujly written C. drprcaiiuf and (fiiprcana. 

1 .MissiM-lt C. 'Iiii>rri^!-(i. 

i:iuoi,o>' HKi.vuM. y 

fig. 1 (1771;: /(/., Hist. Quadr. ii. p. 55:.^ pi. lii. iij,'. 1 (.17eil) ; 

(■(/., op. cif. 3id ed. ii. p. 308, pi. civ. tig. 1 (1793). 
lloiissette jaime, G. Cuvier, Tabl. Hem. cT Ilkt. Nat. p. 104 (1798). 
Wspertilio vaiupyrus (nee L.), var. C, Schreber, Siiug. i. p. lo4 

(1774). Var. y, Gmelin, Limi. St/st. Nat. ed. 13, i. p. 45 (1788). 

Var. 4, Turton, Linn. Syst. Nat. i. p. 24 (180i>). 
Pteropiis vainpvrus, var. y, Er.vlehen, 8i/st. Heyn. An., Mamm. p. 133 

(1777). Var. y, Donndorff, Zo<jl. Berjtr. i. p. 6l> (1792). Var. A, 

Bechstein, Pennant'.^ Uehers. vicrf. Th. ii. pp. G19, 733, pi. liv. fif,^ 3 

(1800). llli(je>; Abh. Akad. Berlin, 1804-11, p. 78, cf. p. 84 

"\'espertilio vaiupynis belviis, Kcir, Animal Ktnqdotn, i. pt. i. 

pp. .wii, 91, no.' 108 (1792). 
PtL>roovon lielvus, K. Ayiderscn, Ann. ^- May. N. II. (7) xix. p. 504 

Pieviipu.s strainintHis, Ji. Genffvuy, Cat. Mamm. Mas. Nation. d'Hist. 
Nat. p. 48 (1803: habitat uuknown) ; id., Ann. Mus. d'Hist. 
Nat. XV. p. 95 (1810 : Tini'ir, errore) ; G. Fischer, Zooyn(ma,\\\. 
p. 557 (1814) ; Oken, Le/irb. Naturyesch. iii. Abtli. ii. p. 9.34 
(1810); G. Olivier, Beqne An. i. p. 124, footuote (1817); 
Desmarest, Nouv. Did. d'Hist. Nat. xxix. p. 512 (1819); id., 
Enrycl. Mcth., Mamm. i. p. 110, no. 143 (1S20); Temminck, 
Mon. Mamm. i. p. 195, pi. xv. %s. 12-13 (skull) (1825) ; Lesson, 
Man. Mamm. p. 112, no. 291 (1827); Gray, in Griffith's 
Aniin. Kinyd. v. ]). 57, no. 100(1827); Desmarest, Diet. Sci. 
Nut. xlvi. p. 366 (1827) ; Is. Geofroy, Diet. Class. d'Hist. Nat. 
^iv. p. 702 (1828); J. B. Fischer, Si/n. Mamm. p. 86, no. 15 
(1829); Wayler, Sy.«t. d. Arnphibien, p. 9 (1830); Lesson, 
Hist. Nat. Mamm. (Coinpl. Biiffon) v. p. 55 (1836) ; Temminck, 
Mon. Mamm. ii. p. 84 (1837: Sennaar, Senegal); Gray, 
May. Zool. ^- Bot. ii. p. 503 (1838); Blainville, Osteoyr. 
Mamm. i. livr. 5, p. 100, At his, Cheiropt. pi. vi. fi<^. 2 (skull) 
(1840: .Sennaar); Wai/ner. Schreber' s Siiiiy., Suppl. i. p. 357 
(1840); Lesson, N. Tabl. lieyne A??., p. 14, no. 188 (1842); 
Siindevall, K. Sr. Vet.-Akad. Handl. 1842, pp. 198, 206 
(1843: Sennaar); Sehinz, Verz. Siiuy. i. p. 129 (1844); 
Temminck, Esq. Zoul. p. 54 (1853: Gold Coast); Wayrier, 
Schreber' s Siiuy., Snppl. v. p. 603 (1853-55); Gervais, Hist. 
Nat. Mamm. i. ]). 189 (1854) ; Giebel, Sduy. p. 999 (1855) ; 
Heiiylin, Beise N.O.-Afrika, ii. p. 14 (1877). 

Xanthai'pyia straniinea, Grai/, List Mamm. B. M. p. 38 
(1843); Gerrard, Cat. Bones Mamm. B. M. p. 58 (1862); 
Fitzingcr, SB. Akad. Wien, liv. Abth. i. no. 10, p. 544 (1866: 
Sennaar, 13abr el Abiad, Bahr el Asrak, Kordofan) ; id., op. cit. 
Ix. Abth. i. Heft 9, p. 458 (1870) ; Gray, Cat. Moi.k. ^c. p. 116 
(1870); liochebnine, Faiine Scueyambie. pt. 2, p. 40* (1883); 
Mafschie, Sdvy. D. Ost-Afrikas, p. 17 (1895) ; Thomas, P. Z. S. 
1896, p. 790 (Zoniba, Nya.<a) ; id., P. Z. S. 1897, p. 927 
(Mt. Malosa, Nyasa); Mafschie, Meyuchiroptera, p. 62 (1899); 
Cliyny, Miss. Shicyal, p. 292t (1900 : Saint Louis, Senegal). 

I'achysonia straiuinpum, Tomes, P. Z. S. 1S()0, p. 44. 

Oynonycteris straminea t, Peters, MB. Akad. Berlin, 1867, p. 866 

* Misspelt Xantarpya straminea. 
+ Misspelt Xauiarpia straminea. 

X VaHousiy written Cyuonyctcris ov Cyioniclcris : C. straiiiiiiiiis. stromuiea, 
ijl' ft r>i mi II ill. 


(tVoiu Sfiuiaar aud Abyssinia to Guinea) ; id., v. d. Decken's 
Reisen, iii. 1, Siiitg. p. 5 (1869) ; Greef, SB. Ges. Naturw. 
Marburg, 1884, p. 44 (San Thoni^) ; Martinez y Saez, An. Soc. 
JEspan. Hist. Nat. xv. p. 339 ( 1 88fi : Fernanda Po) ; Jentink, 
Notes Ley den JiJiis. x. p ■5'2 (1887 : Liberia) ; Monticelli, Ann. 
Mus. Civ. Gifnom, (2) v. p. 524 (pt.) (1887 : Somaliland ) ; Jentink, 
Cat. Osieol. Mamm. p. 264 (1887) ; id., Cat. Synt. Mamm.T^. 152 
(1888); H. Allen, Proc. Ac. Nat. Sci. Philad. 1889, p. 336 
(wing-membranes) ; Bocaye, Jorn. Sci. Math. Lisboa, (2) no. 1, 
p. 15 (1889 : Eio Cuillo, Caconda) ; Bidtikofer, Beisebilder aus 
Liberia, ii. pp. 362, 471 (1890) ; Bre/im, Tierleben, 3 ed. i. p. 349 
(1890: habits) ; Bocaye, Jorn. Sci. Math. Lisboa, (2) no. 7, p. 173, 
fig. (palate-ridges) (1892) ; Matschie, Mitth. deutsch. Schutzyeb. 
■vi. Heft 3, p. 7 (1893: Liberia, Gaboon); Bocaye, Jam. Sci. 
Math. Lisboa, (2) iv. no. 13, p. 4 (1895: Fernando Po, note on 
food) ; Pousaryues, Ann. Sci. Nat., Zool. iii. p. 256 (1896 : 
French Congo)'; Sjo^-tedt, Bih. K. Sv. Vet.-Akad. Handl. xxiii. 
Afd. iv. no. 1, p. 46 (1897 : Camerocn) ; Trouessart, Cat. Mamm. 
i. p. 85 (1897) ; Bocaye, Jorn. Sci. Math. Lisboa, (2) v. no. 19, 
p. 137, fig. (palate-ridges) (1898: Angola, S. Thom^); Seabra, 
ibid. pp. 169, 169, pi, i. fig. 13 (palate-ridges) (1898) ; Socage, 
op. cit. (2) vii. no. 25, pp. 27, 46, 55 (1903: Fernando Po, 
Principe I., Annobon) ; id., op. cit. (2) vii. no. 26, p. 66 

(1904: S. Thomd); Scubra, ibid. p. 103 (1904: Liicinda, 
Pterocyon stramineus, Peters, Jorn. Sci. Math. Li.'fboa, iii. no. 10, 
p. 123 (1871 : Ajuda) ; id., MB. Akad. Berlin, 1876, p. 474 

(Cameroon, Cap Lopez) ; Miller, Fam. ^- Gen. Bats, p. 55, fig. 7 

(auditory buUie) (1907 : Liberia). 
Cynonycteris (Pteropus) stramineus, Marchi, Atti Soc. Ital. Sci. 

Nat". XV. p. 517 (1872-73) (structure of hairs). 
Cvnopterus (Xantbarpyia) straminea, Trouessart, Rev. S{ May. Zool. 

"(3) vi. p. 206 (1878). 
Rousettus* stramineus, W. L. Sclater, Mamm. S. Africa, ii. p. 109 

(1901: Namaqualand, Mashonaland) ; Thomas, in H. H. Johnston's 

The Uganda Protectorate, i. p. 422 (1902) ; Thoma.s, P. Z. S. 

1903, 1. p. 295 (Khartoum); id., P. Z. S. 1904, ii. p. 187 

(Fernando Po ; sexual colour-difference) ; Trouessart, Cat. 

Ma7)im., Siip2)l. p. 59 (1904) ; H. H. Johnston, Liberia, ii. p. 690 

Rousettus (Pteropus) stramineus, Anderson ^- de Winton, Zool. 

Egypt, Mamm. p. 91 (1902 : Upper Nile). 
Yespertilio caninus {nee Blmnenb.), var. b, Goldfuss, Veryl. Natur- 

beschr. Sauy. p. 98 (1809). 
Pterocyon paleaceus, Peters, MB. Akad. Berlin, 1861, p. 423 

Pteropus moUipilosus, IZ". ^//ew, Proc. Ac. Nat. Sci. Philad. 1861, 

p. 159 (1862 : Gaboon). 
Pteropus palmarum, Heuglin, Leopoldina, Heft v. nos. 3-4, p. 34 

(June 1865: Middle and Upper Nile); id., Reise N.O.-Afrika, 

ii. p. 15 (1877). 
Xantljarpyia palniarum. Fitzinger, SB. Akad. Wien, Ix. Abth. i. 

H. 9, p. 456 (1870). 
Xantharpvia leucomelas, Wagner in Hit. {fide Fitzinger) ; Fitz- 
inger, SB. Akad. Wien, liv. Abth. i. H. 10, p. 544 (1866: Sennaar, 

* Variously spelt 7iV?(^'f/')'«s or Eotissettits. , ,, .,•,.. 


Jiiilir el Abi;id, I'.alir el Asrak, Kordol'aii) ; ul., op. cit. Ix. Abtli. i. 

H. 9, p. 401 (1870). 
Leiponyx biittikoferi, Jentink, Notes Leyden Mus. iii. p. 59 (Jnn. 

1881": Liberia); id., op. cit. x. p. 53 (1887: Liberia); id., 

Cat. Syst. Mamvi. p. 152 (1888: Liberia); BiHtikofer, 

bilder am Liberia, ii. pp. 362, 471 (1890) ; Trouessart, Cat. 

Mamm. i. p. 89 (1897) ; Matschie, Meyachiroptcra, p. 85 (1899) ; 

Troziessart, Cat. Mamm., Stippl. p. 04 (1904). 
Pterocyon biittikoferi, Miller, Fum. Sj Gen. Bats, p. 56 (1907). 

The characters of this species, as compared with E. dupreanwa 
and sabceum, are pointed out on pp. 8 & 15. 

Fur. — Fur on head, back, breast, and belly short, closely adpressed, 
on neck longer, more woolly. Face in front of and below eyes almost 
naked. Ears naked posteriorly except at base. Above, humerus, 
proximal two thii-ds or three fourths of forearm, femur, and tibia 
clothed with short, adpressed hair ; interfemoral hairy in the 
middle and for a broad space along the tibiae, extreme lateral 
margin naked ; u})perside of feet short-haired. Below, humerus, 
and proximal half of forearm, membrane along inner and outer 
side of forearm, lateral membrane next to body, femur, proximal 
half of tibia, interfemoral in the middle and along sides of tibia3, 

Colour. — General aspect : grizzled straw-yellow and hair-brown 
above, with a more or less pronounced tawny half-collar in adults. 
Sexual difference : females lighter than males, half-collar paler. 

Adult male — Base of hairs of tipperside almost buff, broad ti])s 
hair-brown ; the colour of the hair-tips predominant, giving the 
Avliole of the upperside a darker aspect than in females, darkest on 
the head, pO'^terior part of back, and upperside of tibia?. Halt- 
collar tawny, brighter than in females. Breast and belly yellowish- 
buff on sides, washed with hair-brown in the middle. 

Adult female — Base of hairs of upperside buff, tips hair-brown ; 
the colour of the hair-bases predominant, giving the whole of the 
upperside a lighter, more buffy aspect, grizzled with hair-brown 
on the crown, middle and posterior part of hack, and ujtperside of 
tibia\ Half-collar less pronounced, rarely brighter than ochraceous 
buff. Breast and belly buff on sides, grizzled with hair-brown in 
the middle. 

^'oung and immature specimens are rather similar in colour to 
adult males, but generally of a somewhat darker shade, and with 
no tawny colour on the neck. 

Se.vual difftrence in size. — Inconsiderable ; in 15 fully adult 
males the length of the forearm ranges from 117"5 to 129 mm. 
(average 122-2), in 14 fully adult females from 119 to 131 
(average 124-7). 

Measurements. On p. 16. 

Ramie. Africa : from Somaliland, Sennaar, and Penegarabia in 
the north, to Nyasaland, Mashonaland, and Namaqualand in the 
sou til. 

Tiipc probalily not in existence. 


VespertUlo vainjjtjrLits Jielvus, Kerr ; 1792. — The species of Eido- 
lon described above was well known to the earh' post-Linneari 
systematists, who put it dowu as a variety of VespertUio (or 
Pterojnis) vamptjrus, L. The earliest recognizable figure aud 
description appear to he those given by Pennant, in 1771 (I. s. c), 
under the name " Lesser Ternate Bat," so called because Pennant 
considered it a lesser variet)' of Seba's " Canis volans Ternatanus 
orientalis." Kerr's V. vampyrus Jielvus is based on Pennant's 
description and figure of this bat. Type originallj' in Museum 
Leverianum ; no habitat given by Pennant, nor by Kerr. Senegal 
may be fixed as the type loculity of E. helvmn. 

Pteropus stramineus, Geoff. ; 1803. — Eased on three examples in 
the collection of the Paris Museum, viz., two specimens (nos. 92 and 
93) from unknown locality, presented by Professor Fourcroy, and one 
(no.94)"de la collection du Stathouder," this latter wrongly supposed 
by Geoffrey to be the original of Seba's description and figure of 
Canis volans Ternatanus orientalis (Thes. i. pp. 91-92, pi. Ivii. 
figs. 1, 2). None of the specimens now in the collection of the Paris 
Museum can bo pointed out, with certainty, as the true cotypes of 
Ft. stramineus. Four characters in Geoffrey's description of this bat 
are, however, decisive evidence that his specimens were E. helvum, 
viz., upper incisors '■ tres-ecartees " ; a short tail (10 mm.) ; total 
length 150 mm., expanse 640 mm.; colour '• jaunatre."— The 
" Cat. Mamm. Mus. Xation. d'Hist. Nat." (1803) was suppressed by 
Geoffroy himself, and the name Pteropus stramineus is therefore 
usually dated from his well-known paper in Ann. Mus. d'Hist. Nat. 
XV. (1810). In this latter Geoffroy based Pt. stramineus on two 
specimens, the one stated to be from Timor (Peron and Lesueur), the 
other without details (and possibly one of the cotypes from 1803). 
The error as to the locality of the former of these specimens was 
pointed out by Temminck (1837, I. s.c). 

Pterocyon paleace^i,s. Pet.; 1861. — Type locality, Africa ; no type. 
Owing to Geofl^roy's statement that Pteropms stramineus had been 
obtained in Timor by ]\^ron and Lesueur, and evidently unaware of 
the fact that this error had been corrected by Temminck, Peters 
proposed for the African species the name Ptcrocifon paleaceus. 
Eut Pt. stramineus was in reality not from Timor, but from Africa; 
and the very species named by Peters ijaleaceus. 

Pteropus mollipiilosus, H. Allen ; 1862. — Type locality : " W. 
Africa"; as belonging to DuChaillu's collections, the type is no 
doubt from Gaboon ; type presumably in the Philadelphia Museum. 
The essence of the original description is this : a very small tail 
present ; first upper premolar larger than incisors ; third digit 8". 
Eidolon hdviihi is the only known Afiican bat for which the 
combination of these characters holds good. 

Pteropus pahnarum, Heugl. ; 1865. — Type locality: "Am mitt- 
lern und oberu weissen Nil und zwischen Senar und Fazogl Ijings 
des blauen Flusses"; type not in existence (?) (see Heuglin, Eeise 
in N.O.-.\frika. ii. p. 15). DiflTers, according to Heuglin, from 
Pt. f/ranuif'is ■" durch Mangel eines Sohwanzrurjiment? luid mit 


Ausnahmo dcr Basis gauz frcieii Daumen."' The pollex is in 
E. helvum always included in the membrane only by the metacarpal 
aud the extreme base of the first phalanx. That the specimen had 
no tail is no doubt a mistake ; if, when skinning an Eidolon, the 
tail -vert ebrnc are extracted together with the body, the empty tail- 
skin shrinks so raiTch that the specimen seems to be tailless. 

Xanlharpyia leucomeJas, Fitz. ; 1866. — In Schreber's ' Saug- 
thiere,' Suppl. i. p. 358, footnote (1840), Wagner describes a 
female of '■^ FUropus stramineus" (presumably in the Munich 
Museum) " aus den oberen Xilgegenden." Even if Wagner had 
not recorded the sex of this specimen, it would be easy to see, from 
the description of the colour of the fur, that it is a female of 
E. helinim. Fitzinger, being unaware of the sexual colour differ- 
ence in this species, ])roposed for the specimen referred to by 
Wagner the name X. leucomelas. A skin in the British Museum 
(specimen c in the list below) belongs to the same series as the type 
of X. leucomelas. 

Leiponyx huttUcoferi, Jent. ; 1881. — Type locality: St. Paul's 
River, Miilsburg, Liberia; type in the Leydeu Museum. Chief 
characters, according to Jentink : postcanine teeth ^; second digit 
■without claw. — I have examined the t3'pe in the Leydcn Museum, 
and find it in every respect indistinguishable from E. helvum, nor 
can I see any difference between British Museum specimens obtained 
at places near the type locality of Z. hiittlkqferi (Nigeria, Ashantee, 
Dahomey)and specimens from other places of Africa; the considerablo 
amount of individual variation in this species, in external dimensions 
and in the size of the skull ai;d teeth, is well shown in a British 
Museum series of fifteen adult individuals fiom Fernando Po. — The 
teeth in the type of L. hiittikoferi are excessively worn, some of 
the posterior molars entirely lost and their alveoli filled out. This 
explains Jentink's statement that the number of cheek-teeth is -., a 
result evidently arrived at as follows : — Upper jaw, left side : p\ p* 
((hese premolars entire), two roots of p^ (m' lost and alveolus closed ; 
of m* a rudiment of posterior root present, but no doubt undetectable 
when the skull was in situ), giving an apparent total of four teeth ; 
upper jaw, right side : p', p^ two roots of p'' (ra' and m" entirelj' 
disappeared and alveoli closed), giving similarly a total of four : 
mandible, left side : p,, p.^, p^ (these premolars entire), a broad 
interspace representing raj (disappeai'ed, alveolus closed), two roots 
of m., (m, entirely wanting), giving seemingly a total of six; 
mandible, right side : p^, p.,. p^, two roots of m,. anterior root of m^ 
(])osterior root of m„ and m^ lost), giving similarly a total of six. — 
As to the absence of the claws of the second digits, it must be said 
that not the claws only but the w'hole ungual (third) phalanx is 
wanting ; the distal articular surface of the second jihalanx is. 
however, in both wings laid bare, nakedly projecting, so that the 
missing phalanges have undoubtedly been violently torn off. The 
I'rcsh condition of the wounds shows that this must have been 
done shortly before, or peihaps after, the death of the individual. 




^ ad. sk. ; 

c? ad. skull. 

2 ad. sk. ; 

(J imm. al. ; 


•2 5 ad. sks. ; 
skull of e. 

Khartoum, Sudan ; Surg. -Major H. N. 

15 Aug. 1902. Dunn [P.]. 

Khartoum, Sudan. Surg.-Miijor H. N. 

Dunn [P.]. 
Sennaar. Purchased(Parreys). 47.5.27-28. 

(Eepresents Xantharpyia leiicomelai, Fitz.) 
Ituri forest, betw. Ruwenzori Explo- 
ration Comm. 

g. Ad. skull. 


(J ad. al. 
3 S ad., 1 2 

ad. sks. ; 

S jun- al. ; 


sks.: skulls. 

(} ad. sk. 

Mawambi and 

Avakubi, 2500' 

{R. E. Dent). 
Ugaya I., Victoria 

jS'yanza, 3000' ; 

30 Dec. 1901. 
Ugaya I., Victoria 

Nyanza, 3000'. 
Mt. Malosa, Nvasa, 

5500'; Nov. 1898 

(.4. Whyte). 
Zomba,Nyasa ; Jan. 

1896 {A. Wliyte). 
Zomba, Nyasa ; 1 1 

Mukimvika, mouth 

of Congo E. ; 

Jan. 1892. 

R. J. Cuniiighame, 
Esq. [P.]. 

R. J. Cuninghame, 2.7-5.3. 

Esq, [P.]. 
Sir J. Kirk [0.]. 68..3.1(>.13. 
Sir H. n. John- 

ston [P.]. 

Sir H. H. John- 

ston [P.]. 
General Manning 


J jun. al. ; 


1 2 imm. 

sks. ; skulls. 

1 d pull, 
sks. ; skulls. 


Juv. sk. ; 

cJ ad., 2 ad- 
sks. ; skulls. 

(5 ad. sk. ; 

Rev. J. M. 


H. Ansell, 

Fernando Po Com- 
mittee [P.]. 



Elloby District, 

Bilelipi, Fei-nando 

Po, 10 m.; 20 

Feb. 1904 (£".&!- 

Bantabiri,Fernando Fei-nando Po Com- 

Po, 10 m.; Feb.- mittee [P.]. 

Mar. 1904 (£•.&/- 

Bantabiri,Feruando Fernando Po Com- 

Po, 10 m. {E. mittee [P.]. 

rellaVista,Principe Hon.W.Rothscbild 

I. ; June 1901 

{A. Mocqiicrys). 


al. ; skull 

of ^'. 
6 ad., S 

jun. sks. ; 
2 ad. al. 

i^. cJ ad. sk. 

San Thome ; Nov. 

Roca Laura, San 

Thome ; April 

1901 {A. Moc- 

Old Calabar. 

Abuchi, S. Nigeria, 
140 m. ; 19 Nov. 

Asaba, S. Nigeria. 


A. Mo(;querys[C.]. 


A. Murray, Esq. 

A. J. Braham, Esq. 

70.3.2iM4-15. 2. 

Dr. W. H. Crosse 

[C. & P.]. 

Expedition [P.], 



Jim. sk. ; 

Wlivdali, Dahomey. 

L. Fi-aser [C.]. 


cJ Hd. sk.; 

Wh\ dab, Dahomey. 

L. Fi-aser [C.]. 


<S ad. ek. ; 




1300' ; 2 Mav, 

fai-d [P.]. 

(S ad. sk. ; 

Kumasi, Ashantee ; 



1 .July, 1898, 

fard [P.]. 

Jiin. sk. ; 


G. Eendall, Esq. 



(J iiuiii. sk. ; 



2 juv. sks. ; 


Purchased (Par- 



Ad. sk. ; skull. 

Not reg. 

3. Eidolon sabaeum, K. And. 

Cvnonvcteris stiauiinea (nee Geoff.), Monticelli, Ann. Mus. Ck\ 
'Gen'oca, (2) v. p. 524 (pt.) (1887 : Yemen) ; Matschie, SB. Ges. 

naturf. Fr. Berlin, 1893, p. 20 (Lahej ; note on habits). 
Xantharpyia straniinea, Yevhiiry Sf Thomas, P. Z. S. 1895, p. 545 

(Lahej ; note on habits) ; Matschie, Megachiroptera, p. 62 (pt.) 

(1899: Lahej). 
Eousettus stramiueus, Thomas, P. Z. S. 1900, p. 98 (Lahej ; note on 

Pterocvi'n sabiBus, K. Andersen, Ann. Sf May. JN". II. (7) xix. p. 505 

(1907, June 1 : Lahej). 

Differs from E. heluiim in the following particulars : — tSkuU 
smaller: total length (one male ad., six females ad.) ol-5-55 mm., 
against 5-t"0-62'2 in E. helvum (nineteen males ad., twelve females 
ad.); maxillary tooth-row (c-m^) 19-2-20-S, against 21-23-5; 
cranial rostrum slenderer ; posterior premolar and molars, above 
and below, markedly broader than in the larger-skuUed E. helvum. 
The external dimensions average slightly smaller. 

Other characters as in E. helvum. 

Measurements. On p. 16. 

Banff e. S. Arabia (Lahej, Aden). 

T)jpc in collection. 

Lahej, Aden. 

Labej, Aden ; o Mar. 

a-b. cJad., (^imm. 

c-f. S ad., 2 ad.,; skulls. 1895. 
/.; skull. Lahej, Aden ; 27 Nov. 

g. $; skull. Labej, Aden ; 19 Aug. 

h-j. (5juv.,22ad. Lahej, Adeu ; 19 Aug. 
sks.; skulls. 1899. 

Col. J. W. Terbury 

[C. & P.]. 
Col. J. W. i'ei-bury 

f C & P 1 
Royal Society [P.]. 99.3. 1 4. 1. 

W. Dodson [C.]. 

(Type of species.) 
\V. Dodson [0.]., 





E.dupreamrm.' JS. hdvum. 


E. snh(P7im. 

^fiN. Max. MiN. Max. Mix. Max. 


Foreanu 1-7 

Poll ex, c. u 49 

2nd digit, metacarpal S8 

„ 1st phalanx ! 17 

2iid-3rd pbalaiix, c. u i l-^'S 

3i'd digit, metacarpal ' 88 

„ 1st phalanx | «^>9 

, , 2nd phalanx 91 

4th digit, metacarpal ; 84 

1st phalanx ' 47 

2nd phalanx j 54 

5th digit, metacarpal I 79"8 

,, 1st phalanx j 37 

2nd phalanx j 335 

Ear, length from notch 

,, greatest width, flattened 1 

Front of eje to tip of muzzle 

Tail ... 

Lower leg . . . 

Foot, c. u ; 39 

Skull, total length to front of premax. ...} 68 

,, width of brain-case at zygomata...' 22 

., zygomatic width 328 

,, width acro>s m-, externally j Ifi 

„ width across c, externally 11'7 

,, palation to incisive foramina ' 28-2 

„ front of orbit to tijD of nasals I 21'7 

Mandible, length 45-2 

Upper teeth, c— ni^ 21'2 

Lower teeth, c — m, 24 

93 5 










91 -.T 


46 8 



































26 8 
















Cyuoiiycicris (pf.), Dob.soii. Cat. Chir. B. M. p. 70. 

1821. Rou.«ettui?, Graxi, London Medical liepository, 

XV. p. 299 (Apr. 1, 1821) R. fe?yptiacn!=>. 

1829. Cercopteiopus, Bnrneft, Qvati. Jmirn. Sci. 
Lit. Art, 1829, pt, 1, p. 269 (Apr.-.Tune, 

1829) I^- PRg-yptiacus. 

1843. Eleuthenna, C?;Y/y,i?s!^ Mamm. B. .If.p. xix. 
Nonien luiduni. 

1843. Xanthaq)Yia, Grau, Lii<t Mointu. B. M. 

pp. xix, 37 I"'- amplexicaunatu.* 

1844. Eleutiier'iira, &'>Y/?/, Vvyafie ' Sulp/mr,' i.T^.29. R. leachi. 
1852. Gvnonycteris, Peters. Beise Mossa7nb., Zool. i. 

'Satiff. p. 25 R- lecachi. 

1870. Senonvcteri?, Gray, Ca>. .Monh. S^x: p. IM. . K. seminiulus. 


Diagnosis. — Geueral shape of skull as in Eidolon, but no bony 
auditory meatus, premaxillaries in contact or co-ossitied in i'ront. 
Incisors f — H ; cheek-teeth j or ^ ; lower incisors (when unworn ) 

2 J ' b b ' ^ ' 

bifid ; p' subequal in bulk to an upper incisor ; ra^ shorter than m, 
and m^ combined. Second digit clawed ; membranes from sides of 
back ; a short tail. Forearm 69"5-99 mm. 

Skull (figs. 2-4 *). — Brain-case in most species deflected very 
nearly to the same degree as in Eidolon, the alveolar line projected 
backward passing through upper part of occipital condyle or upper 

Fig.-. — RousiUus wifirpiiaius, rj. . C;uro, Egypt. r!Cu. 3. 12.S. 1. j. 

margin of foramen magnum ; in one species ( 7i'. lanosus: subgonu-s 
Stenonycteris') the deflection is so strong that the prolongation of the 
alveolar line goes through the middle of the supraoccipital, in another 
{R. anfjolensis : subgenus Lissomjcleiis) so slight that the liiie 
passes below, or througli the lower margin of, the occipital condyle ; 
in all species the deflection is distinctly greater in immatuv© and 
young than in fully adult individuals. Tympanic never elongated 
e.xternally into a bony auditory meatus. Ilostrum comparatively 

Fig. y oil p. 49, fij.'. 4 tiu p. b'l. 


a little shorter than iu Eidolon; from front of orbit to tip of 
nasals less than length of maxillary tooth-row (except in E. lanosus, 
owing to reduction of cheek-teeth in this species), but much greater 
than lachrymal width of skull ; front of orbit vertically above 
middle or posterior half of m^ (cf. Myonycteris). Nasals not pro- 
duced quite as far forward as front of premaxiUaries, Premaxil- 
laries in perfect contact in front, in one species {R. angolensis) 
co-08sified without any trace of the original suture. Postorbital 
processes not reaching halfway between frontal and zygoma. 
Sagittal crest low or undeveloped, the temporal ridges often remain- 
ing separate throughout the life of the individual. 
Other cranial characters as in EidoJon. 

,, i' i' c p' p' p^ m' m'' ^ o . 

Dentition (figs. 2-4 *),— Normally v-. 5— LJ X 2:= 34. 

^ iii.cp, P3P, m^m^m, 

p^ and m^, p, and m^ reduced ; p^ permanent in most species, 

deciduous in li. seniinudns and brachyotis. A minute m^ occasionally 

present on one side or both sides t. In one skull in the collection 

(R. anr/olcnsis, S ad. ; G.12.4.5) a small supernumerary molar 

(very broad, but excessively compressed antero-posteriorly) is 

present on the right side, closely wedged in between m' and m'^. 

Upper incisors equidistant, or i'-i' slightly more separated than 
i^-i^. Upper canine and p' generally widely separated, sometimes 
rather closely approximated (individually in R. ampler icaudatus 
and, particularly, R. brachyotis), very rarely in actual contact 
(occasionally in R. brachyotis when p' is wanting), p' much more 
reduced than in Eidolon, being in cross section only equal to (or 
smaller than) an upper incisor, in some species deciduous (R. s'^ml- 
■nudus, brachyotis). m^ much smaller than ra', but not reduced to 
the same degree as in Eidolon. 

Lower incisors subequal in height and bulk, or the outer ones 
slightly larger in cross section ; cutting-edge in youngish individuals 
distinctly bifid ; the median notch of the cutting-edge continued 
for a short distance down the front of the crown as a faint vertical 
groove ; this groove is usually detectable even -when the cutting- 
edge has been worn straight, p^ small, in most species from twice 
to four times the bulk of a lower incisor, in one species {R. ango- 
lensis) not, or scarcely, exceeding a lower incisor ; as a rule situated 
nearer to the canine than to p^. m^ nearly always exceeding 
(sometimes only equalling) p^ in length, but never quite as long as 
m and m^ combined, m, smaller than, sometimes only half the 
size of, m^. m^ much smaller than m.^, elliptical or subcircular in 

The molariform teeth, above and below, are uuusually narrow in 
R. lanosus ; an approximation to this is found in R. celebensis. 

* Fig. 3 on p. 49, fig. 4 on p. 52. 

t Details from 88 skulls examined, representing all species known : — 
m' present on one side : R. leacAi, cue adult, teeth well worn ( ; 
/?. aewimtdiis, one adult, teeth unwora (nut registered) ; m-* present on both 
sidf s : R. (pgijptiacuii, one adult, teeth practically unworn ( 


ruhtti'-riihjes. — Slightly varying according to species and indi- 
viduals: — (1) In most species {11. Imchi, arahicis (soinctimes), 
leschenavlti, semini(diis, anijjJexicaii.ilatns, celebensix) 4 + 3-)- 1 ; first, 
ridge terminating at p\ or between c and p' ; second at p^ ; third 
at front of p^ ; fourth at front of m', or betAveen p^ and m' ; fifth 
afc m", or between m^ and nv ; sixth more or less closely behind m' ; 
seventh far behind m" ; eighth situated at palation border. Some- 
times the fourth ridge is interrupted in the median line, the formula 
being 3-}-4-f-l ; this I have seen in B. lanosus, IL hraclnjotis, and 
some individuals of li. leachi, and according to Bocage it is 
(normally ?) the case in R. angolensis. A ninth, more or less 
indistinct, ridge is occasionally detectable (formula 4 + 3-1-2, or 
3 + 4 + 2 ; see R. angolensis). (2) In some species (R. agj/ptiacus, 
often in arahicvs, rarel}- in amplfxicau(fatic) an additional divided 
ridge is developed behind the sixth; formula 4 + 4 + 1 ; but the 
extremities of this ridge are either connected with, or situated 
closely behind, those of tlie sixth ridge. 

Head. — Muzzle proportiotiall}' somewhat shorter than in Eidolon ; 
measured from front of eye to tip of nostrils shorter than (only 
in R. lanosus almost equal to) distance from front of eye to 
base of ear. Nostrils separated by a deep groove ; inner margins 
moderately projecting. Inner margins of li])S Iringed with short 
pa])il]8e. Median portion of tongue with oblique lows of rounded 
papilla?, each papilla with three small, straight, sharply pointed, 
backwardly directed horny spines. 

Ears. — Tip of ear-conch reaching hinder corner of eye : outer 
much more convex than inner margin, tip rounded ; in some species 
(7^. arabicu-t, angolensis. htnosus) a shallow, but distinct, emargina- 
tion of outer margin below tip of ear. Antitragal lobe small, flatly 
rounded (most sjiecies) or triangular (R. angolensis), in one species 
(7^. hinosus) practically wanting. 

Wings. — Chief characters : second digit clawed ; membranes 
from sides of dorsum and back of first or second toe. 

I'irst phalanx of first digit one and a half the length of meta- 
carpal. Second digit subequal to (a little longer or shorter than) 
third metacarpal. Third metacarpal from six to seven tenths the 
length of the forearm, a little longer than fourth metacarpal, which 
is a little longer than fifth. First ])halanx of third digit two third? 
to tlircc fourths the length of the metacarpal ;■ second jihalanx 
four fifths to six sevenths of metacarpal. First jihalanx of fourth 
digit rather more than htdf of metacarpal : secotid phslunx a little 
longer than first. First phalanx of fifth digit equal to, or a Tutle 
less than, half of metacarpal; second phalanx a little longer or 
shorter than first. See wing-indices, on p. 2<l. 

First digit included in the raenibrano by the metacarpal and base 
of first phalanx. 9-15 long fascia' in the lateral membrane, 
viz. 2-4 jMistaiiconeal, 7-13 preanconeal. 

Tail. — From less than one half to five sixths the length of the 
hind foot ; basal portion connected with interfemoral by its dorsal 
integument, terminal portion fr-'clv projecting. 

c 2 





2nd digit. 

3rd digit. 

4th digit. 

5th digit, j 





























B. leachi, espi/pfiacvs, arabicus . 

JR. lesckenaulH, seminudus, ^ 
anipfericaudaivs, hrachyotis j 




^oot. — Tarso-metatarsal portion of foot, long : measured on 
plantar surface equal to or more than two thirds the length of 
phalanges with claw of third toe (compare Myonycteris). 

Pur. — In niosit species: fur ou bod}- short, adprcssed; face in 
front of and below ejes very short-haired ; notopatagium naked ; 
upperside of tibiae short-haired or naked. In three species (ll.cele- 
bensis, avgolmsis, and lanosus) the coat is longer, velvet or (lanosus) 
rather coarse ; face more distinctly haired ; notopatagium and 
upperside of tibiae well haired. In all species the fur of the body 
extends ou the proximal half or two thirds of the forearm. 

Sexual differentiation. — Inconsiderable. Glandular hairs on fore- 
neck and sides of neck more brightly coloured in adult males of 
certain species {E. angolensis, ample xicaudatus). Females seem to 
average a trifle larger than males, but the difference, if any, is 
infinitesimal. In some species (e. g. 11. leachi and lesc/ienaulti), of 
which the majority of individuals are dark-coloured, specimens 
occur which have the upperside more or less suffused with Prout's 
brown or mars-brown, the underparts with wood-brown ; the 
available material is not extensive enough to show v\hether this 
colour-difference is sexual, dependent on age, or indicative of the 
existence of two colour-phases. 

Range. — The African continent, exclusive of the Mediterranean 
countries AV. of Egypt ; S. Asia, from Palestine and Cyprus to 
S. China; the Indo- and Austro-Malayan Archipelagos, as far east 
as the Solomon Islands. 

Habits.- — The fruit of £'rzo6oi/'2/rt ("Loquat') is the favourite food 
of the S. African ii. leachi; when eating the bat throws itself on to 
a neighbouring branch or su.spends itself on quivering wing.s, and 
seizing the fruit in its mouth either bites a portion of it away at 
once or pulls it off from its hold ; in this way it destroys far more 
frait than it eats ; in default of fruit it devours in.';ects, snapping 
them off the flow'ers and leaves without alighting. (Layard.) 

In Egypt, H. (Fgyptiams is found in ancient tombs and temples, 
old mosques, and Sheiks' graves, or in crevices of rocks ; in the 
days of Etienne Geoffroy it frequented the recesses of the Great 
Pyramid, hut now (Dr. J. Anderson writes, in 1902) it is not found 
in that building ; it is also met with in trees in gardens, and inj 

date-plaiitatious around villajrcs, iu sycamore, mulberry, and other 
trees; occasionally it is found singly, but generally a few are 
associated togeilier ; feeds on fruits of the date-palm, wild figs, &c. 
Visiting Cyprus in 1901-2, iliss D. M. A. Bate found li. (pr^i/ptiacas 
excessively common ; it does considerable damage to the fruit-crops, 
particularly to the oranges and dates, though while the latter are 
ripening they arc often envelo])ed in sacks or matting for protec- 
tion ; during the summer it roosts in thick trees, in winter in 
closely packed bunches in the roofs of old buildings and caves ; it 
is very restless, and extremely noisy, even in daytime and when 
undisturbed (P. Z. 8. 1903, ii. p. 342). 

Ji. leacM has repeatedly bred in captivit)- (Zoological Gardens, 
London and Cologne). During the act of copulation the male is 
suspended (as usual by thn hind feet) behind the female, its under- 
parts being iti contact with the lower back of the female; an 
embracing with the anterior extremities does not take place ; 
period of gestation (in one case, Cologne gardens) precisely 1.5 weeks 
after last copulation ; during the first months after birth the young 
bat was rarely seen, suspended as it was from the mammfe of the 
female and completely covered by her wing-membranes ; by tlie 
end of the third month the young commenced to take part in the 
meals of the parents (the juice, not the flesh, of moist fruit), but 
remained in its protected po-itiou at the breast of the mother till 
the end of the fourth month ; at the age of eight months it was 
still not quite full-grown (Wundcrlich, Zool. Garten, xxxii. pp. 78- 
81, 1891). — According to Dr. J. Anderson, j-oung of R. cer/)/j)tiacHS 
are born generally in February and Jlarch ; a female of M. arabicns 
killed on March 29 at Lahej, Aden, had a single young one at her 
breast, and other gravid females had single foetuses (Yerbury & 
Thomas, P. Z. S. 1895, p. 545); a female of ii'. amphxicmdatus 
obtained by A. Everett in Alor Inland, N. of Timor, had one young 
on March 24 (collector's label, British Museum). 

Scarcely anything is known about the habits of the Oriental and 
Austro-Malayan species. Hodgson's account of the habits and 
extensive niglitly migrations of " Fferojms pyrivorus" i. e. B. le- 
xcJienanlfi, in Nepal (J. A. S. B. iv. p. 700. 1835) — copied by Hutton, 
P. Z. S. 1872, ])p. 693-94, under the head of Cynopterus marginatiis, 
and by Dobson, in his Catalogue, under Cynonycteris amplexi- 
caudatvs, and again, by an oversight, under Cynopterus marginaf^'s 
— is based wholly on a misconception, as pointed out by Scully 
(J. A. S. B. Ivi. pp. 237-38, 1887). Dobson was informed that a 
colony of 7^. lesrhi'navlti living near the sea at ifoulniein were seen 
to feed on Mollusca left exposed by the tide (J. A. K. B. x)ii. p. 200, 
1873) ; this would seem to explain the occurrence of an allied 
species (7?. arahicns) in so dreary a desert as the island of Kishm, 
Persian Gulf, where an exclusively fruit-eating mammal could 
hardly exist. 

Affinities. — Rousettus is allied (though not very closely) to 
EiloJ.on ; the two genera represent no doubt diverging branches 
from one common stem. In having no bony auditory meatus, the 


jjreinaxilhiries in coiir.>icL (not separated), lu, not lengthened, and lU" 
less reduced in size, Rousettus is less specialized tlian Eidolon ; but it 
is on a higher level in the rather shorter rostrum and more reduced p\ 
The range of the genus over the whole of the Ethiopian and Oriental 
regions, the close affinity of ]l. arahicus to the S. African 2i. Uachi, 
the absence of any representative of the genus from the whole of 
the Srediterraneau subregion * (except Egypt), are evidence that its 
origin dates back to a time when, owing to different physiographic 
conditions, Africa andS. x\sia were much more intimately connected 
than now. Eidolon is a peculiarly modified Ethiopian offshoot of 
the common prototype. 

The eleven known species fall into three natural, rather sharply 
separated groups (subgenera) : — 

(!) Subgenus Eousettus : — Brain-case moderately deflected ; 
premaxillaries in contact, not co-ossified (except sometimes in 
li. mjiiptiacm) ; cheek-teeth unmodified in size and shape ; p^ 
much larger in bulk than a lower incisor; wings from first toe; 
antitragal lobe small, but distinct. — This is the least specialized 
of the three subgenera of Housettus, in all essential characters 
perhaps the most primitive group of living Vlegacbiroptera. Range : 
the Ethiopian Itegion, through Egypt, Cyprus, Palestine, Arabia, 
8. Asia, the Indo-Malayan and Austro-Malayan Archipelagos, 
eastward to the Solomon Islands. The nine species are referable 
to three types : — (a) K. lecichi, (egi/j^tiaai^, and arahicus : rather 
heavily built species, with strong rostrum and teeth, the second 
phalanx of third digit lengthened, the poUex comparatively long : 
distributed over Africa generally, Cyi)ru'5. Palestine, Syria, and 
Arabia, as far as Sind (Karachi), li. ivi/i/pticicns is a larger-skulled 
modification of the li. leachi type ; li. arahicua is more closely 
allied to the S. African R. leachi than to R. agyptiacm. — (b) R. le- 
scheiiaidti, seminadus, nnrple.vicaudatus, wiinor, and hracliyotis : very 
closely related to the species of the former section, but rather more 
delicately built, wirh slenderer rostrum, feebler teeth, the second 
phalanx of tiie third digit not lengthened, the pollex comparatively 
shorter. The members of this section are probably on the whole 
slightly less specialized than those of the former. R. lexdiemmlti 
(continental S. Asia) and semiuudxs (Ceylon) come near to the 
S. African Ji. leachi in the width of the interspace between c and p^, 
the size and shape of m^, the width of the ears, and the shortness 
ot the tail ; in the Indo-lfalayan R. ampleA-icandatus there is a 
tendency to a reduction of the diastema c-p\ m^ is smaller and 
more circular in outline, the ears narrower, the tail averaging 
longer, the general dimeiisions smaller ; most of these characters 

* Botisettus fft/i'ltiri/i. 'froupssart ("Cat. Miuiiui., Suppl. p. fiO, 1904; baped 
on Cf/iioni/cteris (?) sp., Claude Gaillard, C. R. Aonrl. Sci. cxxv. p. (i20, 1897, 
and Arch. Mus. d'Hist. Nat. I,yon. vii., 2 meiu. p. 6, fig-. 1, 1899), known from 
a complete right humerus and the dista! extieinities of a risfht and lelt humerus, 
from the Middle Miooene of La Grive Saint-Allian, Lsere. is not a liuuseftus, 
nor even a Fruit- Br,t, but some large species of Microchiroptera, as proved by 
Gaillard's figure of the humerus (1. s. c. : high, flange-like deltoid crest). 

EousErius. 23 

find a climax in the Austio-Malayau li. brachyotis : diastema c-p'^ 
still more reduced, p' deciduous, ears still smaller, size smaller. — 
(c) li. celebensis : palate and cheek-teeth narrower than usual, fur 
louf^er and richer, size small, wings proportionately long; probably 
a modification of the R. amplcxicaudatus type. 

(2) SxENOXiCTEUis* (subg. nov.): — Brain-case strongly deflected ; 
preraaxillaries never co-ossified ; check-teeth excessively narrow : 
width of p' about one fifth (in llousettus s. str. about one third) 
that of palate between fronts of p' ; p^ much larger in bulk 
than a lower incisor; wings from second toe; antitragal lobe 
obsolete; fur long and rather coarse. Etliiopian. One species: 
R. lanosns, Thos. — So far as the reduction of the cheek-teeth is 
concerned, this peculiar species stands in the same relation to the 
rest of the genus Rouset/iis as the narrow-toothed species of Pleropus 
(Ft. sahniger, personatus, woodfordi, sc(i2ndatus) to the normal- 
toothed Fteropi. If its habits were known, R. lanosus would 
probably prove to subsist cliiefly on food (juice of fruits) tliat 
requires little or no mastication. 

(3) LissoNTCTKRis t (subg. nov.) : — Brain-case only slightly 
deflected ; prcmaxillaries co-ossified in front, even in }'oung adults ; 
cheek-teeth peculiarly short and broad, subquadrate ; p^ reduced 
in size, being only subequal to a lower incisor ; wings from second 
toe ; antitragal lobe distinct ; fur limg and silky. Ethiopian. 
One species : R. aw/olensis, Boc. — This is the most aberrant species 
of the genus ; although in all important respects a " Rousettus," it 
shows, in the small deflection of the bi'ain-case, the shape of the 
occipital region of the skull, and the outline of the cheek-teeth, 
leanings towards the genus Epomophorns. 

Rousetius, Gray ; 1821. — Xo description. Type (only species): 
Pta-opus orpiptiacus, Geoff. The name Ronsctfii.s must have been 
suppressed or forgotten by Gray ; it does not occur in any of his 
later jtapers. Revived by Palmer, Proc. Biol. Soc. Wash. xii. p. 112 
(30 April, 1898). 

Cercopteropus, Burnett ; 1829. — No description. Two species 
mentioned : C. (e.fjifpliacus and C. amplexicandatus. The former, as 
being the first-named, may bo fixed as the type. 

XantJunpi/ia, Gray; 1813. — Xo description. Three species, 
enumerated in the following order: The Xantharpye, Xantharptfia 
ample.vimndata ; Egyptian Xantharpye, X. wgyptiaca ; The Pale 
Xantharpye, A", stniminea. Both on the " first species principle " 
and on the " tautology principle " A'. ampJe.vicaudata must be re- 
garded as the type of the genus. It should be mentioned that the 
two examples referred by Gray in this book (List Mamra. B. M. 
p. 37, 1843) to "X. ample-v'icavdata" are in reality not this 
species but Dohxonia pnllala ; however, by his reference to Geoffrey's 
original description and figure of Pteropus atnplexicaiidatus Gray 

* Irevos (narrow), ri'icrfpis (bat) ; in allusion to the I'xcessively narrow 

t AiiT(TOf (soft), rvKTfpis ; from the silkv character of the fur. 

2i KousKrrus. 

clearly iudic-ittes wliicli species is meant bj- the name ample.vicau- 
(lata. — A brief diagDOsis of the genus Xanthnrpiiia was given by 
Gray in Voyage of the ' Sulphur,' i. p. 29 {1844): distinguished 
from Eleutherurn by having the " tail with the base inclosed in 
the underside of the interfcmoral meml^rane." 

Eleutheriira, Grray; 1844. — First occurrence of the name in Gray's 
List Marara. B. M. p. xix (1843): no description, no species. 
Second occurrence in Voyage of the ' Sulphur,' i. p. 29 (1844) : 
type (only species mentioned), E. hnUentotta, Temm. ; a brief 
diagnosis showing that Gray based the genus on Temminck's 
erroneous description of Ptero'pus hottentottus : tail free (not con- 
nected with interfemoral), projecting from " a nick on the middle 
of the narrow interfemoral"; but Temminck's Pi. hoUeniotius is 
A. Smith's /'t. Jearln. 

Ci/noiii/cteris, Peters ; 1852. — Type (only species mentioned) : 
Pieropus coUnris, Licht. (nee ill.), i. e. Pteropiis Icachi, A. Smith. — 
The genus Cjfnoru/cterh, as understood by Peters in 1867 (Kevision 
of Megachiroptera, MB. Akad. Berlin, 18G7, ])p. 86.5-866), included 
all the then known species of liousettus and Eidolon. Dobson's 
Ci/nonj/ctens (Cat. Chir. 1878) corresponds to the genera Eidolon, 
liousettus, and Mi/oni/cteris of this Catalogue. 

Senoni/cteris, Gray ; 1870. — A subgenus of Xantlmrpiiia. Typo 
(only species) : Xantharpyia scminuda, Gray. Separated from 
Xantharpijia s. str. on account of its "very sparse and s'hort " 
fur, "especially on the back," and " nearly bald" forearm. 

tSi/nopsis of the Species.^ 

J. Brain-case conspicuously detJectfd ; preujax- 

illiiries in contact ; p, much larger in bulk 

tlian a lower incisor. 

A. Brain-case moderately deflected ; width 

of p^ about one tliird tliat of palate 

between fronts of p^ : wings from 1st toe ; 

antitragal lobe distinct Subg. lioisETTUs. 

a. Pollex .'iO-37-.5 : 2nd phalanx of third 

digit 50-5-61-5 mm. 

«'. l<;ars not attenuated at tip ; lower leg 

40-4 So mm. 

«-. Length of skull 40'-'j-1.'>-8 mm. ; 

rostrum slenderer : palate-ridges 

normally 4-f 34-1 : forearm 89- 

99 mm.' (S. Africa) }. H. k-achi. p. 2.5. 

b'. Jjeni;th of skull 4o-6-4()-7 nun. : 
rostrum heavier ; palate-ridges 
4 4-44-1; foreanii 88-99 mm. 

(Angola to Palestiue) i'. R. (/■i/i//)f;,irf/s, y. -J'.}. 

I'. Ears attenuated at tip ; lower leg .'17- 
*•"■ •'j9-5 mm. ; forearm 87-96. (Arabia 

to Sind) y. Ji. unihieu.-: p. 3-1. 

h. P.dlex 24-30; 2nd phalanx of 3rd digit 
••;(;- 17-2 mm. 

" On n. ,»/,/.>;■, sve p, 4;l a.ud Acldeiidt at rnd of Voi.ii.i 

I'.uusrrms lkacui. 

c . Miihirt^ not iiiuisually narrow ; fur 

sliort: Botopatag'iuin naked. 

cK m., elliptical in outline, once and a 

half or Iwice as long' as Lioad ; 

width of ears 14'0-15'8 mm. 

ff\ p' not deciduous; fur on nape 

and shoulders not unusually 

scarce ; for'earm 80'5-87'5 mm. 

(India, Himalaya, to S. China) 

b^. pMeciduous; nape and shoulder.^ 

seuiinaked ; forearm 79-85/5 

mm. (Ceylon) 

d'. m.. iu outline, only 

slightly longer than broad ; width 

of ears 10-1. 3 mm. 

<■". p' not deciduous ; forearm 77- 

87'2 mm. (Indo-Malayan) .. 

cP. p' deciduous; fosearm 69-8-76 

mm. (Austro-Malayan) . . . . 

d'. ^Molars unusually narrow ; bony 

palate narrow ; fur longer ; noto- 

patagium and tibife well haired ; 

forearm 72-5-75 mm. (Celebes) . . 

B. Brain-case strongly deflected ; cheek-teeth 

excessivelv narrow ; width of p' about 

one Kfth tLat of jialate between frimts of 

])* : wings from :^ud toe ; antitragal lobe 

1 'bsolete 

c. Fur long and coarse ; lower leg 39-40 ; 

forearm 88-5-90 mm. (Ethiopian) . . 

11. Brain-case slightly deflected : prema.xillaries 

co-os?ifiod ; pi subequal in bulk to a lower 

incisor ; molars short and broad : wings 

from 2nd toe ; autitragal lobe distinct . . 

d. Fur long and silky; lower leg 29-31 ; 

forearm 79-83-5 mm. (Ethiopian) . . 

[p. 35. 
4. li. Ifschenaulh, 

5. R. seminuJus, p. 3S. 

LP- -lO. 

6. 1{. amplcxkaudatus, 

8. R. iruchyotis, p. 44. 
9. R. celcbensis, p. 4G. 


10. R. hmosus, p. 49. 

Subg. LissoxYCTEnis. 

11. R. angoknsis, p. 51. 

1. Rousettus leachi, A. Sm. 
Cffuonycteris cuUaris, Dobson, Cat. Chir. B. M. p. 75. 

I'teropus collaris {nee III.), Licit rtisfei'n, J'erz. Douhl. Mas. Berlin, 
p. 3, no. 47, p. 5 (1828 : Terra Caffrorum) ; Giehel, Sdug. p. 1000 

Cynonycteris collaris, Pctcvf:, Rrise MoRsambiqne, Sdiif/. p. 25 (1852: 
liiliaiiibane); id.. MB. Akad. Bfirlin, 1867, p. 865 (S. Africa); 
r. L. t'idnfp); P. Z. S. 1868, p. 404 (at sea oil" Natal); ojx cit. 
1869, p. 602 (Natal) ; op. cit. 1870, p. 127 (fig. of $ ad. with 
young) ; op. cit. 1871, p. 478 (breeding in captivity); op. cit. 
1873, p. 193 (specimens in captivity); Dohson, Cat. Chir. Ind. 
Mv.-<. p. 2 (1874 : S. Africa) ; Gulliver, R. Z. S. 1875, p. 493, 
pi. iv. tig. 7 (red blood-corpuscles) ; Dobson, Mo>i. A.<iat. Chir. 
p. 190 (1876: S. Africa) : J. AiidprsoTi, Cat. Mamm. Lid. Mm. 
1 p. 104 (1881 : S. Africa) : Jeidink, Cat. OstroK Mamm. 
p. 263 (1887: Cape); id., Cat. Si/af. Mamm. p. 151 (1888); 
//. Al/cii, Proc. Acad. Nat. Sci. Philad. 1889, p. 337 (wing- 
membranes) ; Brehm, Tierlebi->i. 3 ed. i. p. 350 (1890) ; Wunder- 
lich, '/.oil. (Uirtcn. .\.\xii. i)o. .'>. p. 78, toxt-fig. (I'^Ol : breeding in 
ciplivily t : 'I'miiv^mrl . Cat. Mmtmi. i. p. 8J ipt.) (1^07). 



Cvnopterus colLiris, Kolenati, Moii. europ, Chir. p. 11 (18(30: Cape, 

Eleutherura coUaris, Gray, Cat. Monk. ^-e. p. 118 (1S70: Cape, 

Cynopterus (Cynonycteris) ccllaris, T^-ovessart, Jiev. ^- Mag. Zool. 
_ (3) vi. p. 206 (pt.) (1878: Cape of G. H., Natal). 

Xantbarpyia collaris, Matschie, Megachiroj^tera, p. 66 (pt.) (1899). 

Koiisettus collaiis. W. L. Sc/afer, 3famm. S. Afr. ii. p. 106, 
text-fig. 118 (1901 : Cape, Natal) ; Anderson ^ de Winton, Zool. 
Egypt, Mamm. pp. 86, 88 (1902) ; Troitessart, Cat. Mamm., 
Sup2)l. p. 60 (pt.) (1904) ; Thomas ^- Schcann, P. Z. S. 1906, 
p. 161 (Knysiia) ; Mtller, Fum. Sf Gen. Bats, p. 54 (1907). 

Pteropus am])lexicaudatus {nee Geoff.), Temminck, Man. Mamm. 
i. p. 260 (1827 : Cape) ; J. B. Fischer, Syn. Mamin. p. 86 (pt.) 
(1829: Cape). 

Cynouycteris amploxicaudata, Dohson, Cat. Chir, B. M. p. 73, 
.specimens e-j (1878). 

Pteropus leacbi, A. Smith, Zool. Journ. iv. p. 433 (1829: Cape); 
id., Bidl. Sci. Nat. xviii. p. 272 (1829 : Cape) ; Temminck, in 
Smuts s Enum. Mamm. Capens. p. 5 (1832: Cape); A. Smith, 
S. Afr. Quart. Joiiryi. ii. p. 53 (1833 : Cape) ; Lesson, Hist. Nat. 
31a77im. [Compl. Buffon) \. p. 49 (1836: Cape); Temmijick, 
Mon. Mamm. ii. p. 88 (1837: Cane); Gray, Mag. Zool. 
^ Bot. ii. p. 603 (1838 : Cape) ; tVagner, Schreber's Sdug., 
Suppl. i. p. 361 (1840 : Cape) ; Biippell, Mas. Senck. iii. Heft 2, 
p. 154 (1842: S. Africa); Lesson, N. Tabl. Eigne An. p. 14, 
uo. 192 (1842 : Cape) ; Schinz, Syst. Verz. Sdug. i. p. 131 (1844 : 
Cape) ; A. Smith, 111. Zool. S. Afr. pi. 48 (wbole fig., bead, skidl) 
(1847: Cape); Wagner, Schreber's Sdug., Suj)pl. v. p. 604 
(1853-55: Cape, Mozambique); Gervais, Hist. Nat. Mamm. i. 
p. 191 * (1854) ; Schlegel, Eierkunde, i. p. 63 (1857 : S. Africa) ; 
Luyard, Cat. S. Afr. Mus., Mamm. p. 19 (1861). 

Xantbarpyia leacbi, Gerrard, Cat. Bones Mamm. B. M. p. 57 (1862 : 
Natal) ; Fitzi^iger, SB. Akad. Wien, Ix. Abtb. i. Heft 8, p. 466 
(1869: Cape, Mozambique). 

RoLisettus leacbi, K. Andersen, Ami. ^ Mag. N. H. (7) xix. p. 506 
(19071 ; Lonnberg, Kiliimmdjaro-Meru E.vpedilion, pt. 2, p. 6 
^908: Tanga). 

Pteropus hottentottusf, Tejuminck, in Smuts's Enum. MammXkqjens. 
p. 3 (1832 : Cape) ; id., Mon. Mamm. ii. p. 87, pl. xxxvi. figs. 16, 
17 (skull) (1837: Cape); Wagner, Schrebei'^s Sdug., Suppl. i. 
p. 360 (1840: Cape) ; Eilppell, Miis. Senck. iii. Heft 2, p. 154 
(1842: S. Africa) ; Lesson, N. Tabl. Rh/ne An. p. 14, no. 191 
(1842 : Cape) ; Schinz, Syst. Verz. Sdug.'i. p. 131 (1844 : Cape) ; 
Wagner, Schreber's Sdug., Suppl. v. p. 604 (1853-55 : Cape) ; 
Gervais, Hist. Nat. Mamm. i. p. 191 (1854 : S. Africa) ; Giebel, 
Sduf/.-p. 1000 (1855); Grid, IC Sv. Vet.-Akad. Handl. n. s. ii. 
no. 10, pp. 9, 13 (1860: Knysua). 

Xantbarpyia bottentotta, Blyth, Cat. Mamm. Mus. As. Soc. p. 21, 
no. 56 71863: S. Africa); Fitzinger, SB. Akad. Wien, Ix. 
Abtb. i. Heft 8, p. 468 (1869 : Cape). 

Cynopterus brevicaudatua [nee Is. Geojf.), Gray, List Mamm. B. M. 
p. 39 (1843). 

Kousettus sjo.stedti, Lonnberg, Kilimandjaro-Meru Expedition, pt. 2, 
p. 7 (1908: Tanga) t. 

* 'Misspelt Pteropus ErachU. 

1 Earliest spelling : Jwt/cntof/uf ; niifspeU hoflenfoUis by a fen- of the 
juUiurs quoted. \ 8ee Addeui.x 



/>;(r/(K»s«'s.— Distiu^niished from all other species of the genus by 
the combination of the following characters:— Frontal region of 
skull between postorbital processes flattened; premaxillaries in 
co)itact, but rarely co-ossiiied ; total length of skull 40-5-43-8 mm. ; 
palate-ridges normally 4 + 3 + 1. Wings from back of first toe, 
or interspace between first and second toe; pollex (with claw) 
31 -35-5 mm.; second phalanx of third digit 50-5-60 mm. ; second 
phalanx of fifth digit nearly always shorter than first plialanx ; 
cars not attenuated at tip ; fur short. Forearm 89-99 mm. 

Skull. — Brain-case moderately deflected (compare E. lanosus and 
Ji. cmgnlensis). rremaxillaries in contact in front, hut rarely, 
even iii very aged individuals, co-ossified (compare li. (pgijptiacus 
and R. nnr/oleiisis). Frontal region between postorbital processes 
flattened (compare R. anr/olensis) ; region between orbits cora- 
))aratirely narrow, the width of the interorbital constriction in 
fully adult specimens being almost always distinctly less than the 
width of the postorbital constriction (compare R. cef/i/ptiacus). 
Temporal crests uniting into a sagittal crest at level with root of 
zygomata, but very often remaining separate throughout the life of 
the individual. 

Teeth. — Upper incisors almost eiiuidistant, or the interspace 
between i' and i' much less than between i* and c. Distance c-p' 
equal to or larger than between i" and c. p^ small, equal in size 
to i-. m' equalling or exceeding p' in length, m' rather less than 
half the size of m\ — Lower incisors crowded. A broad interspace 
between c and p,. p, in cross section from twice to four times the 
size of a lower incisor, situated nearer to ihe canine than to p^. 
m . m,, and ra , elliptical oval or almost elliptical in outline ; m.^ 
somewhat smaller than m, ; m^ from two thirds to less than one 
half the si/e of m.,. 

J'alate-ridgex.—'SoTmaih 4 + 3-|-l. Variations from the rule: 
the fourth ridge in some individuals interrujited in the median 
line (formula: 3 + 4 + 1); a 9lh, more or less indistinct, ridge 
occasionally detectable (formula : 4 + 3+2). 

Ears. — Outer much more convex than inner margin; tip not 
attenuated, broadly rounded off above (compare R. arabicus). 
Antitragal lobe flatly rounded. 

Winqs. — Pollex with claw equal in length to four fifths of second 
metacarpal (or equal to first phalanx of fourth digit). Second 
digit equal to metacarpal of third digit. First phalanx of third 
digit eijual to two tliirds of metacarpal; second plialanx only a 
little shorter than metacarpal. Second phalanx of fifth digit, with 
rare individual exceptions, shorter than first phalanx (compare 
R. avr/olensis). Wing-indices, see anted, p. 20.— Notopatagium 
semicircular in outline, naked ; lateral membrane inserted on end 
of first metal arsal, or between first and second toe. 10-14 long 
f.isciic in the lateral membrane, viz.. 2-3 postanconeal, 8-10 pre- 

7'(/(7. — From one half to three fourths the length of the hind 


/.',„.._l.;;irs naked posteriorly, except at base. Face in front of 
and below ev.s short-haired. " Ftir on back short, soft ; longer. 



more wooll)', and less closely adpressed on foreneck. Fur of body 
extending upon upperside of humerus and proximal half or two 
thirds of forearm ; front margin of antebrachial membrane fringed 
with short hairs ; notopatagium naked ; femur well haired ; tibia 
and hind foot to claws clothed with very short and thinly spread 
hairs ; central portion of interfemoral well haired, lateral portion 
along tibiae almost naked. Belowj proximal two thirds of forearm, 
plagiopatagium next to body, femur, and central portion of inter- 
femoral covered with short, thinly spread, woolly hair. 

Colour.— General aspect : above, uniform dark brown with a 
tinge of slate, or more or less suffused with Prout's brown ; below, 
uniform smoke-grey, or more or less suffused with wood-brown. 

Eack and rump varying from dark hair-brown tinged with slate 
to bistre ; sides of back and rump next to membranes, in adult 
specimens of both sexes, often approaching Prout's brown, this 
colour sometimes extendiug over the whole of the upperside of the 
body ; crown and occiput markedly darker than back ; nape of neck 
lighter hair-brown. Underside uniform grizzled smoke-grey; in 
adult specimens of both sexes the foreneck, flanks, and under 
surface of humerus and forearm are often more or less suffused 
with wood-brown. / 

Measurements. On p. 34. 

Range. Cape Colony, Natal, Lower Zambesi (Inhambane), north 
to German East Africa (Tanga). 

Gotijpe in collection. 

Pteropus collaris, 111. ; 1815. — Type locality : " die ostlichen 
[afrikanischen] Inseln." — llliger's Pteropus collaris (Abh. Akad. 
Eerlin, 1804-11, pp. 71, 84; published 1815) is Erisson's 
" Roussette a col rouge" (1756"), Euffon's "Eougette" (1763), 
Kerr's Vespertilio vampi/rus subnir/er (1792), E. GeofFroy's 
Pteropus ruhricoJlis (\S10). In 1823 Lichtenstein (l.s.c.) wrongly 
identified llliger's Pt. collaris with the S. African F'ruit-Bat here 
under consideration, but the error, hidden as it was in the little- 
known" Verz. Donbl. Mus. Berlin," passed for manj' years unnoticed, 
the species being constantly referred to as Pteropus leachi or Pt. 
liottentottns. In 1852 Peters (' Peise nach Mossambique ') confirmed 
Lichtcnstein's wrong identification of Pt. collaris, and from about 
that year the names Imchi and hottentotttts gradually went out of 
fashion, being replaced by collaris : from about 1870 leachi and 
hottentottns only appear in the lists of synonyms of collaris. 

Pteropus leachi, A. Sm. ; 1829. — Type locality : " Gardens about 
Cape Town"; cotype in collection. — Summary of description: 
" supra fusco-cinereus, infra sordido-cinereus, cauda libera." The 
earliest available technical name of the species. 

Pteropus hottaitottus, Temra. ; 1832. — Type locality : " circa 
urbem Capensem "; type in the Leydeu Museum (authentic specimens 
in the British Museum, see list below, specimens a-e). — Original 
description contributed by Temminck to Smuts's ' Enuraeratio 
Mammalium Capensium.' Temminck points out the differences 
between Pt. hottenfofius and Pt. amj-ih.ricyjtrlafvs, but was admittedly 
in doubt whether the former was separable from Pt, leachi 


('■ de Fterofio leacJn tenendum vidptiir, eiim forfasse a praecedente 
[i.e. Ft. hottnitotto] nou distinctum esse specie"). Owing to 
a passage in the original description ("ad basin caudae perbrevis 
eernitur incisura, figurara inversae V pi'ae se ferens ; juxta banc 
Cauda, quae plane est libera, exoritur "), Ft. hottentottns As-as by 
subsequent writers ((iray, Waguer, and others) supposed to differ 
from Ft. leacJii by a V-sliaped eraargination of the interfemoral at 
the base of the tail, and by having the tail perfectly free from the 
interfemoral, diB'ercnces which undoubtedly were due to shrinkage 
of the median part of tlie interfemoral in the mounted, type of 

a-e. 4 ad., 1 iimn. ]^ear Cape Town Purchased (Leyden, 
sk.s.; skulls. (J.Sinuls). Museum). 38,40,67. 

{ Autbentie specimens of I'teropus hottentoiius, Temm.) 

f-h. 2 S "d-. 1 6 Cfipe Town. Trustees of the S. 

iuv. al.; skull Africaia Museum 

of./; [P.]. 

/. Ad. skull. Cape Town (5i> .J. H. Ford, Esq. [P.]. 


(Figured in 111. Zool. S. Afr.) 
j. $;ul.;il. Cape Town (.Sir .-1. Surg.-Gen. G-. E. 

ISmitli). Dobson [E.]. 

( Cotijpe of species.) 
k. S j"T- -^k- ; Cape of Good Hope. Purchased (Ver- 
skull. reaux). 

l-f. 4cJ'ad..4 $ ad. Knvsna. 3 April, C. 1). Rudd, Esq. 
sks.; skulls. iyuo (C. //. .S. [P.]. 


f-v. (S ad., 2 id., Kny.-5nu {C. H. B. C. D. Eudd, Esq. 
$ juv. al. Grant). [P.]. 

w. Ad. St. ; skull. Natal. Purchased (Stock- f 

holm Museum). 

.r. c^ad. al. Natal. Dr. Seem;ni [C.]. 

y.; skull. S.Africa. Purchased (Brandt). 

2. Rousettus asgyptiacus, E. Geoff. 
Ctjnonycteris cegypiiaca, Dobson, Cat. Chir. B. M. p. 74. 
Pteropus ivgyptiacus •*, E. Geoffroy, Ayin. Mus. d'Hist. Nat. xv. 
p. 90 (ISiO: Lower Egypt); Oken, Lehrh. Natiirgesch. iii. 
Abth. il. p. 934 (1816) ; G. Cuvier, Begne Anim. i. p. 124 
(1817); E. Geoffroy, Descr. de rEgypte, Hist. Xaf. n., Manim. 
p. 134, pi. iii. fit;-. 2 (1818) ; Desmclrcst, Nouv. Diet. d'Hist. Nat. 
xxix. p. 513 (1819 : Cairo) ; id., Encycl. Meth., Maimn. i. 
p. Ill, no. 144 (1820); Gray, Londmi Medical Eqxmtory, xr. 
p. 299 (1821); Lichfemtein, Verz. Douhl. Mus. Berlin, p. 3t 
(1823: Egypt); Lesstm, Man. Mamm. p. 112, no. 292 (1827: 
EgTpt) : Gray, in Griffith's Anim. Kingd. v. p. 57, no. 161 
(1827 : Egypt) ; De-'^, Diet. 8ci. Xat. xlvi. p. 367 (1827 : 
Egypt, " Senegal ") ; J. B. Fischer, Syn. Mamm. pp. 85, 549 
(1829 : Egypt, " W. coa.<t of Africa") ; E. Geoffroy, Hist. Nat. 
Mamm., le?. 13, p. 24 (1829 : EgApt) ; A. Smith, S. Afr. Quart. 
Journ. ii. p. 53 (1833: Egypt); " Waterhouse, Cat. Mamm. 3/j/.s. 
Zool. Soc. 2nd ed. p. 13, 'no. 104 (1838); Gray, Mag. Zool. 

* Spelt eqyptiacus by Geoffroy in 1810 (original description of the spesies, 
/. s. c), m 1818 by the same author (Descr. de I'Egypte, I. s. c.) cegyptiacu^ ; the 
former may tlierefore be considered a slip or mi.sprint corrected by the author 
himself. " t Spelt Ft. crgiipiiii^. 


c^- Bot. ii. p. 503 (1838 : N. & E. Africa) ; BJainviUe, OsL 
Mamm. Atlas, Cheiropt. pi. xiii. tig. 2 (teeth) (1840) ; Gervais, 
Hist. Nat. Mamm. i. p. 191, fig. (teeth) (1854: Egypt, Nubia); 
Giehel, Stint/, p. 999 (1855); id., Odontof/r. p. 9; pi. iv. fig. 3 
(teeth) (1855) ; Unger S)- Kofscki/, Die Insel Cyiiern, p. 570 
(1865) ; Gasco, Yiayqio in Erjitto, pt. ii. p. 95 ( 1 876) ; Klunzinqer, 
Upper Egypt, p. 148 "(1878). 

Xantharpyia fegy])tiaca. Gray, List Mamm. Ii. M. p. 37 (1843 : 
Egypt) ; id., List Osteol. Specim. Ii. 31. p. 10 (1847) ; Kolenati, 
SB. Akad. Wien, xxix. p. 344* fig. 30 (palate-ridges) (1858j ; 
Gerrard, Cat. Bones Mamm. B. M. p. 57 (1802: Egypt) ; Tris- 
tram, P. Z. S. 1866, p. 93 (Palestine) ; Fitzimjer, SB. Akad. 
Wien,\\\\ Abth. i. p. 544 (1866: Egypt) ; id., op. cii. Ix. Abth. i. 
p. 463 (1869 : Egypt, Syria) ; Maischie, Me<iachiropt.era, p. 66 

Pachysoma fegy^ptiacuni. Tomes, P. Z. S. 1860, p. 44. 

Cyno'nycteris segyptiaca, Peters, MB. Akad. Berlin, 1867, p. 865 
(Egypt) ; Dobson, Moti. Asiat. Chir. p. 32 (1876) ; Tristram, 
We'dern Palestine, p. 25 (1884) ; .Tentink, Cat. Osteol. Mamm. 
p. 263 (1887: Egypt); id.. Cat. Syst. Mamm. p. 151 (1888: 
Egypt, " Senegal" j ; //. Allen, Proc. Acad. A'at. Sri. Pliilad. 
1889, p. 337 (wing-niembraiie.=) ; Brehm, Tierleben, Sod. i. p. 350 
(1890: habits); Noack, Jahrb. Hamb. Wiss. Anst. ix. p. 56 
(1891: Egypt); Troucssart, Cat. Mamm. i. p. 84 (pt.) (1897); 
Seabra, J. Sci. Math. Lisbon, ( 2 ) v. no. 19, pp. 158, 1 69 ( 1898 : Syria) . 

Eleiitherura a'gvptiaca, Grai/, Cat. Monk. ■^■c. p. 117 (pt.) (1870: 

Cynopterus (Cvnonycteris) jiegvptiaca, Troaessart, Rev. S,- Mar/. Zool. 
"(3) vi. p. 206 (pt.') (1878 : Egypt, Palestine). 

Rousettus t feg"\ ptiacu;', Anderson ^- de Winto7i, Zool. Eqx/pt, 
Mamm. p. 84, "pi. xv, (whole fiu.) (1902) ; D. M. A. Bate, P. 'z. S. 
3903, ii. p. 341 (Cyprus) ; Trouessart, Cat. Mamm., Suppl. p. 60 
(pt.) (1904) : Seiina, Arch. Zool. Napoli, ii. fasc. 3, p. 256 (1905 : 
Erytlirea) ; K. Andersen, Ann. i§- Maq. N. II. (7) xix. p. 507 
(1907) ; Miller, Earn. ^- Gen. Bats, p. 54 (1907). 

Pteropus geoffroyi, Temminck, Mon. Mamm. i. p. 197 pi. xv. 
figs. 14, 15 (JkuU) (1825: Eg^-pt, ""); Is. Geoff roq, 
Diet. Class. d'Bist. Nat. xiv. p. 702 (1828); Lesson, Hi^t'. Nat. 
Mamtn. (Compl. Bufon) v. p. 50 (1836: Egypt, ''Senegal"); 
Wayner, Schrebers Sduq., Suppl. i. p. 358 (1840: Egypt, 
"Senegal"); Riippel', Mus. Senck. iii. Heft 2, p. 154 (1842: 
Egypt); Lesson, N. 'lahl. Reyne An. p. 14, no. 189 (1842: 
Egypt, "Senegal"); Schinz, Syst. J'erz. Siiug. i. p. 130 (1844: 
Lower Egypt, " Senegal '") ; Waqner, Schreber's Smiq.. Suppl. v. 
p. 603 (1853-55: Egypt, "Senegal"); Hevylin, Reise N.O.- 
Afrika, ii. p. 15 (1877 : Egypt). 

Cynonycteria (Pteropus) geoflroyi, Marchi, Atti Soc. Ital. Sci. Nat. 
XV. p. 517 (1872-73: structure of haii-s). 

Eleutherura unieolor. Gray, Ctd. Monk. S^c. p. 117 (1870: Gaboon). 

Cynonycteris collaris {nee 111.), Dobson, Cat. Chir. B. M. p. 76, speci- 
men a (1878 : Gaboon) ; Giinther, P. Z. S. 1879, p. 741 (Cyprus) ; 
Dobson, Rep. Brit. Assoc. 1880, p. 173 (Cyprus); Trouessart, Ctd. 
Mamm. i. p. 84 (pt.) (1897: Cyprus) ; Bocaqe, Jorn. Sci. Mtith. 
Lisboa, (2) v. no. 19, p. 137 (pt.), text-fig. 5 (1898 : palate- 
ridges) ; Seabra, ibid. pp. 159, 169, pi. i. fig. 14 (1898 : palate- 

* Spelt XfiDtharpyia at/yptiea. 

t A'arioiisly spplt RvuseitUf or HtmsffUn^ by the authors quoted. 

Cynoptenis (Cynomcteris) collaris, Trouessart, Rw. ^- Mnq. Zool. 

(3) vi. p. 206 (pt.) (1878: Gaboon). 
Xantharpvia collaris, Mntschie. Megachiroptera, p. 66 ^pt.) (1899). 
Kousettus collaris, Trouessurt, Cat. Mamm., Suppl. p. 60 (pt.) 

Cynonyctei-is sp., Bocage, Jom. Set. Math. Lisbon, (2) ii. no. 7, 

p. 176, te.\t-tig. 3 (palate-ridges) (1892; Pungo Audongo). 

Diagnosis. — Similar to 21. leavhi, but with larger skull, broader 
rostrum, broader frontal region, and heavier teeth ; palate-ridges 
normally 4-f-4-f-l. Forearm 88-99 mm. 

Skull (fig. 2, on p. 17). — Similar to that of li. leachi, but larger 
(see measurements, p. 34), and with markedly deeper and broader 
rostrum. Premaxillaries often co-ossified. Frontal region between 
orbits comparatively very broad, the width of the interorbital con- 
striction in fully adult specimens distinctly larger than the width of 
the postorbital constriction. Owing to a stronger development of 
the temporal muscle (heavier cheek-teeth), the temporal crests unite 
into a sagittal crest at a short distance behind the postorbital 
processes, a sagittal crest being always developed in mature age 
and often present even in individuals with almost unworn teeth. 
For the same reason the coronoid process of the mandible is higher, 
the angular portion stronger and more projecting. 

Teeth (fig. 2, on p. 17). — As in IL leachi, but averaging markedly 

Palate-n'Jr/es. — 4-(-4-t-l, i.e. essentially as in Ii. leachi, but 
with an additional middle (divided) ridge behind the molars; the 
formula given here is taken from a few alcoholic specimens only ; 
some slight individual variation may be found. 

External characters. — General size averaging a trifle larger than 
that of Ii. leachi, head proportionally much larger, muzzle deeper 
and broader, ears larger, but precisely of the same shape as in 
a. leachi; wing-structure as in H. leachi; upperside of tibia and 
foot, to the claws, more densely haired. 

Colour rather similar to that of It. leachi, but of a somewhat 
lighter shade. Back and rump hair-brown, with a slight tinge of 
brownish slate ; crown of head and occiput darker than back ; 
nape of neck light hair-brown, or drab. Underside grizzled smoky 
grey ; foreneck and flanks in adult specimens often more or less 
suSused with wood-brown. In dried specimens exposed to light 
the slaty tinge of the upperside very soon disappears, the colour 
fading into brown or yellowish brown. 

Measurements. On p. 34. 

lian(je. From Loanda and Gaboon to Egypt, Erythrea, Syria, 
Palestine, and Cyprus. 

Tjipe in the Paris Museum. 

Fterojnis agyptiacus, Geoff. ; 1810. — Type locality : " la basse 
Egypte " (" le plafond d'une dcs chambres de la grande Pyramide ''). 
Based by Geoffrey on " plusieurs individus '* ; of these one onlj- 
ajipears now to be in the Paris Museum, an adult male, mounted, 
in bad condition, much faded, skull in situ ; labelled Egypt (lieg. 
no. A. 69). The species was figured eight years later in 'Descrip- 
tion de I'Egypte ' {I. s. c). 



Pleropus geoffroi/i, Temm. ; 1825. — Name proposed in lieu of 
cegypfiacas, " vu que I'espece se trouve au Senegal, et probablemeufc 
sur toute la cote septcntrionale d'Afrique," 

Eleutherura uiiicolor, Gray ; 1870.— Type locality : Gaboon ; 
type iu collection. — Gray did not give any tangible character by 
which to discriminate E. unicohr from E. rpr/i/ptiacus. The type 
is a faded specimen, indistinguishable in cranial and dental 
characters from examples from Egypt ; a specimen in the British 
Museum from Loanda (Pungo Andongo) is further evidence of the 
occurrence of R. cegypiiacus in W. Africa. 

ReniarJiS. — R. mjypt'mcas is a large-skulled northern represen- 
tative of the R. leaclii type. It has not rarely been confused 
with the allied R. leachl and R. arahkus, more frequentlj' with 
the very different R. lanosus. Its distribution in Africa between 
Angola and Egypt, and the exact limit:^ of its area in Syria and 
Palestine (as compared with that of R. arahicjis), remain to be 


(S ad. fk. ; 

Pungo Andongo, 

Dr. W. J. Ansorge 


Loanda, 1200in. 
9 June, 1903. 

; [0.]. 


5 ad. sk. ; 


Purchased (Ver- 


(T//pe of Eleutherura. unicohr, Gray.) 


Ad. sks.; skulls. 


Dr.Turnbu'.l Christie 
[0. & P.]. 

39 «,^'. 


Ad. skulls. 


Dr.TurubuU Cliristie 
[C. & P.]. 

.39 c,./; 


2 2 ad. sks. ; 


Jas. Burton, Esq. 

39 c. </. 


Imm. sk. ; 

Tomes Coll. (A''i- 





Cairo, Egypt 

Mrs. J. Anderson 


1 foel,. al. : 

{Dr. Innc.x). 


skulls of y, 

I, M. 


2 ad. al. 

Assiut, Egypt. ; 

Dr. J. Anderson 

21 Jan. 1892. [C. & P.]. 
/5 .?. cJ ad., 5 ad. Mehalleh el Dr. J. Anderson [P.]. 
al. Kebir, Egypt 

{G. 11. Kent). 
Wady Kurn, Canon JI. B. Tris- ()'i~4H. 

Acre, Pales- tram [C.]. 

Mt. Lebanon; Saleem Baroody [C.]. 94. .5.7.2. 

March, 1894. 
Cyprus. J^ord Lilford [P.].,.l-(;. 

r-s. 2 (S jiin. al. ; 
skull of r. 

t. 2 ad. sk.; 
u-z. 3c5'ad.,32 ad. 
al. ; skulls of 
II? -V-. c? ad., 2 ^^- Cyprus 

^--f. c?ad.,2c5'jun., 

2 ad- 

2 Jan. sks.; 

skulls of 

Lord Lilford [P.]. 

Nicosia, Cyprus Miss D. M. A. Bate 312.4.1-.3, 
440-700' ; [C.]. 5-6. 

Apr., Oct., 
Nov., 1901-2. 


2 juu. 

Ktema, Cypru?, Miss D. M. A. Bate 
200'; 18 Mar. [C.]. .;'. ■ 




3. Rousettus arabicus, And. ^- dc WInt. 

Cyiionycteris iimploxicaudata {tii'c Genff.), Dobson, Oct. Chir. Ttid. 

Mils. p. i' (pt.) (1874: Kisliin I.): Blanforrl, E. Persui, ii. p. Kl 

(1S70: Kislmi I.); Itoh.vm, Mon. Asiat. Chir. p. 190 (])t.) 

(187(3: ki-ihiii I.); J. Andfrsim, Cat. Mamm. Ind. Mus. i. 

p. 104 (pt.) (1881: Kl.sluiil.): Murmii, Vi-rt. Z<i„l. Sind. \->. :i 

(1S84: Miikklre Hill.-, Siud); Trouessait, Cut. Mamm. i. p. 81 

(pt,)(1897: Kishiu I.). 
Cyiio])terii9 (Cvnoiiycteris) amplexicaiid.itii.'',7Voz^e.s.s«;-/, lirv. Sf Maq. 

^Zool. (3) vi/p. -iC) i (pt.) (1878: Per.-sian Gulf). 
Xanthuipyia auiplexicaiiilata, lilanford, Fnuna Brit. Lid., Mamm. 

pt. ii. p. 2U2 (pt.; (1891; Ki.shiii; Karachi); Thomas, P. Z. S. 

1894, p. 44i) (-MiHcat) ; Matschic, Me/ackiroptera, p. 67 (pt.) 

RousHttii.s amiilexicaudatii.'^, Thomas, P. Z. .S'. 1900, p. 98 (Aden) ; 

Troue.smrt, Cut. Mamm., Viippl. p. 00 (pt.) (1904 : Aden, 

^[ll8cat. Iv.i.slini). 
XaiUiiarpvia aig-\ptiaca [nee Geoff'.), Yerlurij Sj- Thomas, P. Z. 8. 

]S9.'5, p. .•')4o (Aden). 
Xanthai'])via collavis {nee III.), Matschie, Meqaehiroptera, p. 66 

(pt.) (1899: Aden), 
lloiiseitiis arabicus, Anderson ^~ dc Winton, Zoo}.. Ei/ypt, Mamm. 

pp. 8(i, 88, 89-90 (1902: Aden); K. Andersen j Ann. i^- Mai/. 

N. H. (7) xix. p. 507 (1907). 

Diagnosis. — Similar to It. IracM, but with shorter and lower 
ro^itriuu, narrower ear-tips, shorter tibia and foot. Forearm S7- 
96 mm. 

Details. — The skull of R. arabicus averaaes in ever}' respect 
smaller than in R. leaclii: total length in fully adult individuals 
138 7- 41 "8 mm., against 40'o-43'S in R. leachi; rostrum markedly 
shorter and slenderer : length from front of orbit to tip of nasals 
];j-18'6 mm., against 13'8-15'2 in R. leachi. Teeth as in 
R. leachi, but smaller, and molars narrower. Palate- ridges some- 
times 4-f-3-f 1, as in R. leachi, but very ofton there is, behind the 
6th ridge, a more or less distinct additional ridiie, the extremities 
of which are either in direct connection with or terminating very 
closely behind thoseof the 6th ridge, giving the formula 4 -|- 4 -f 1, as 
in R. cpi/i/ptiaciis. 

Inner margin of ear-conch neaidy straight, tip of ear much 
narrower than in R. leachi • tibia and foot noticcabl}' shorter. 
Distribution and colour of fur as in R. ii'i/i/ptiacus. 

From R. o'ljijittiacas this species is readily distinguished bj" its 
much smaller skull, shorter and slenderer muzzle, much narrower 
ear-tips, shorter wdngs, and shorter tibia and foot. 

Measurements. On p. 3-1. 

Range. From Arabia (Aden, Muscat) to Sind (Karachi). 

Ti/pe in collection. 

Remarks. — R. arahicus is more closely related to the S. African 
R. leachi than to its geographical neighbour R. ar/if/diacus. The 
exact limits of its area (to the west, north, and north-east) arc still 
unknown ; it is not unlikely tliat it extends into Syria and 



«-f. 2(5a(l.,2 2a(l., lahej, Aden. Col. J. W. Yerbiiry 95.6.1 o-C. 

1 inill. al. [0. & P.]. 

/■.; skull. Lahej, Aden ; Col. J. W. Yerbiirv 
21 Mar. 1895. [C. & P.]. 

(T>/pe of ^pp^!ies.) 

^-/i. c? ad., 5 imm. Lahej, Aden; Col. J." W. Yerbui-y 95.G.1.48-49. 
.^ks.; skulls. 21 Mar. l.'-9.=i. [C. & P.]. 

J-;. c?ad., c? imm., Lahej, Aden ; W. Dodson [C.]. 

2 ad., 22 Aug. 1899. 
jimiu.sks. ; 


m-7?. c? ad., gimni. Lahej. Aden. Percival & Dodson 
al. [C.]. 

0. 9 iirun. al. Muscat, Oman. Dr. A. S. G. Jayakar 

[C. & P.l. 

p-n. 2 ad., 9 inini. Muscat, Oman. Dr. A. S. G. Jayakar 
al. [C. & P ]. 

r. 9 ad. al. Karachi, Sind. Karachi Museum [E.]. 

Measurements of llonsettus leachi, 11. fpgyptiacus, and E,. arabiciis. 

7?. hachi. E. (pgyptiacus. 11. arahkin 

MiN. Ma.\. 


Pollex, c. u 

2nd digit, metacarpal 

,, 1st phahiiix 

,, 2rd-3rd phalanx, c. u ; 

3rd digit, Dielacarpal 

,, 1st phalanx 

I ,. 2nd phalanx '• 

I 4th digit, metacarpal 

„ 1st phalanx 

,. 2nd phalanx 

5th digit, metacarpal 

,, 1st phalanx t 

„ 2nd phalanx 

Ear, length irom notch s 

,, grpate:.t widtl), flattened 

Front of eje to t'p of muzzle 

Tail j 

Lower leg 

Foot, c. u 

Skull, total length to front of premax — 
,, width of brain-rase at z3goniata...i 

„ zygomatic widtli 

., posto]-bilal constriction 

„ interorbital constriction 

„ width across m-, e.xternally 

„ width across c, externally 

,, palation to incisive foramina 

„ front of orbit to tip of nasals i 

Mandible, length , 

Upfer teeth, c— m^ ! 

Lower teeth, c — m^ 




24 7 



12 5 

19 7 

13 8 

19 2 
18 8 
15 2 

Wi.\-. Max. 





























37 2 
18 8 

MiN. M.\x. 






50 5 
















50 5 





4. Rousettiis leschenaulti, Desm. 

Ci/iionifcteris amfiltxicawlnta (pf'.)> Dob.son, Cat. 
Chir. B. M. p. 12. 

Pteropus lesclieiiiiLilti, T)esmarest, Eiiq/cl. Meth.. Mti/iiin. i. p. ]1(), 
110. 142 (IHiJO: Pondichery) ; ifl, Did. Sci. Nat. xlvi. ]>. 3()o 
(1S27: Pondichery); Lesson, Man. M/inwi..\). 110, no. 28;'. 
(1827: Pondichery); (ri-(u/, in Grfffif/i'.s Aiiivi. Kini/d. v. 
p. 5<i, no. 158 (1827: Pondichery) ; Is.. Geofrai/, Diet'. Class. 
d'Hist. Nat. xiv. p. 702 (1828 : "Pondichery) ; J. Ji. Fischer, 
Syn. Mamm. pp. 8ii, 549 (1829 : Pondichery) ; Lesso7i, Hist. Nar. 
Mainm. { (hmjil. Buff(m) v. p. 47 ( IH'Mi : Pondichery) ; Temvnnck, 
Man. Mamm. ii. p. 86 (1837: Pondichery, Calcutta) ; W<i;iiier, 
Schrelicr's &an(/., Suppl. i. p. .'359 (1840: Pondichery, Calcutta) ; 
Lesson, N. Tuld. llhpie An. p. 14, no. 190 (1842 :' Pondiclierv, 
Calcutta); SMnz, jS>/st. Verz. Savg. i. p. 1,80 (1844: Pondi- 
cliery) ; Wai/iu'i; Sc/ireOer's Sii'icj., Siifpl. v. p. 604 (pt.) (185.3- 
55: Pondichery, Calcutta); Giebel, Simp. p. 999 (pt.) (1855: 
Pondiclicry); Bbjtlt, Vat. Mamm. Mus. As. Soc. p. 21, no. 54 
())t.) (]8(!8: Coromandel ; Jcrdon, Mavnn. Itid. ■^. 19 
(1867: Madras, Trichinopoli) ; Sterndale, Mamm. India, p. 40 
(pt.) (1884: Calcutta, Madras, Pondichery, TrichinopoJi). 

Xantharpvia leschenaulti, Fitzinyer, 8B. A/cad. ll'icn, l.\. Abth. i. 
Heft 8,"p. 472 (pt.) (18()9: India). 

Cynonycteris (Pteropns) lescheuiiulti, Mavchi, Atti Soc. Ital. Sci. 
Nat. XV. p. 517 (1872-7."} : structure of hairs). 

Cynouycteri.s leschenaulti, I'eters, MB. Akad. Berlin, 187.3, p.. 485 
(type of species re examined). 

Itousettus lesclienaulti, K. Andersen, Ann. ^- Maq. N. II. (7) xix. 
p 507 (1907). 

Pteropus ainplexicaudatus {neo Geoff.), Tennninck, Man. Mamm. i. 
p. 200 (pt.) (1825 : Siani) ; Desmarost, Diet. Sri. Nat. xlvi. p. .367 
tpt.) (1827: Siam) ; J. B. Fischer, Si/n. Mamm. pp. 86,549 (pt.) 
(1829: Simn); Giebel, Siiuij. p.lO0O(pt.) (1855: India). 

Cynonycteris amplexicaudata, Peters, MB. Akad. Berlin, 1867, 
p. 865 (pt.) (Penj^'al, 8iani) ; Swinhoe, P. Z. S. 1870, p. (ilO 
(Anioy) ; Peters, P. Z. S. 1871, p. 51:'. (Burma) ; iJohson, Proc. 
A.S.B. 1872, p. 154 (Burma); Macalister, Phil. Trans. 1872, 
p. 125 & .«eq., pi. xiv. fifi-. 4 (myology) ; l)obso)i, J. A. S. B. xlii. 
]it. ii. p. 200, footnote ( 187.3) ; id., op. cit. xlii. pt. ii. p. 202 
(pt.), pi. xiv. iig. 8 (ear) (187.3) : id., Cat. Chir. Ind. Mas. p. 2 
(pt.) (1874: Coromandel coast, Siugbhoom, Peo-u): id.. Mm. 
Asia': C'A(V. oPP- -^•', 1'^^ (j't.) (1876: Indian Pen.) ; ? Leche, 
Lnnd.<: Univ Arsshr. xiv. p. 17, pi. ii. fig-. 9 (1878) (milk-teeth) ; 
./. Ander-'ion, Cat. .Mamm. Ind. Mns. i. p. 103 (pt.) (1881 : Coro- 
mandel coast, Iiidia) ; ScxtUij. J. A. S. B. Ivi. pt. ii. no. ,3, 
y. 237 ( 1SS7 : Nepal) ; Jentink, Cat. Oste<d. Mamm. ]). 263 |p(.) 
(1887: Calcutta); id., Cat. Si/st. Mamm. p. 151 (])t.) (1888: 
Calcutta) ; Troue.<sart, Cut. Mamm. i. p. S4 i])t.) (1897). 

Xantharpvia amplexicaudata, Fifzinyer, SB. Akad. Wiev, Ix. Ahth. i. 
Heft 8' p. 470 (pt.) (1869: .Siam); Blanford, J. A. S. B. 
Ivii. pt. ii. no. 3, p. 271 (1888); id., Faun. Brit. hid.. Mamui. 
pt. ii. p. 2(>1 (pt.), text-iifr. 76 (ear) (1891 : India, Himnlava, 
JSurnui); Thomas, Ann. .Sins. Civ. Genora, (2) x. p. 921 (lS92: 
Moulmcin) ; Mn/svhie, Mei/arhirojitcra, p. 67 (pt.) (1809) ; S. .V 
Flower, P. Z. S. 1900, p. 340 (Lao.^ Mts.). 



L\viii'])toi'us (Cyiii nycteris) ainplcxicandatiis, Troiwssarf, Iter. Sf 

' Maij. Zcol. (?>) vi.'p. I'OG (pt.) (1878: Inciia, TsVptil, Jiiiniia). 
CynonTctt'ris (Pteropus) ani)>lexicaudatus, Thcohnld, in Masons 

JJiirmii, i. ]). 'l2o (1882 : Eenf>-al, S. India, Biiniia). 
lit.iisetUis 1 nipU'.\ic,uidiitu8, Trouessarf, Cat. Mamm., Sitppl. p. 00 

(pt.) (1904: Himalaya). 
PteiM pus [,sp.], IMr/.soii, J. A. S. R. no. 7, p. 340 {\^?,± : Nepal). 
I'teropiis pvri\orii^, Ilodqsoii, J. A. S. B. iv. no. 48, p. 700 (1835: 

Nepal) ; ' id., V. Z. S. imn, p. 4G (Nepal) ; uh, Icon. ined. (B.M. 

copy) pi. xli. fifr. 1 (col. fit;'.), pi. xiii. fig. 2 (col. fig.). 
Pteropus pii-ivarus (sic), Hodyaun, J. A. S. B. x. p. 908 (1841 : 

Oynopteiiis afiinis, Gray, List Mamm. B. M. p. 39 (1843 : 

Cynopterus ninrpnatus {nee Geoff.), Gray, List Mamm. B. M. 

p. 38, speciuieriR A, / (1843 : Nepal, the cotvpe.s ri Pt. pyriwrus, 

Hodgs.) ; Biyth, J. A. S. B. xiii no. l.OO, y. 479 (1844 : 'Ne} al) ; 

Gray, Cat. Hodgson Coll. B. j\J . ]). 3 (184(j : Nepal) ; Ilidton, 

P. Z. S. 1872, p.' (J93 (N.W. Hiiiialaya). 
Eleutlierura niar^inata, Gray, Cat. Blonh. S)-c. p. 118 (1870: 

" Nepal," really Nasirabad). 
'■■: Eleiitlierura fuli<:inosa, Gray, Cat. MovJc. i^c. p. 118 (1870: Laos 

Mts., Siani). 
.: Eleutlierura fusea. Gray, Cat. Monk. S)-c. p. 119 (1870 : 

" India P"). 
Cynonycteiis infuscata, Peters, MB. Akad. Berlin, 1873, p. 487 


Diagnosis. — Allied to ii. arahicvs, but f-maller, with tlie imizzle 
shorter and slenderer, tlie tip of the ears not attenuated, the jiollex 
markedly shorter, wings shfirter, cs]jecial]y the first and second 
phalanx of the third digit, and the loot smaller. Porcaim 80-5- 
87"5 mm. 

^ill•uU and teeth. — Similar to those of Ji. arahicvs, but skull 
averaging a little smaller, rostium slendeier; preniaxillaries ratlier 
more strongly projecting forward, palalion border more t'harjily 
angulate. Teeth on the whole ratlier smaller, raolais somewhat 
narrower ; p' not deciduous, unless perhaps in very aged individuals 
(cf. li. femini(tlns) ; m^ quite or almost equalling m, in length ; 
m, elliptical in outline (cf. 2i. umph x.cuudaiits). 

Fulate-ridges. — 4 + 3 -+- 1 ; number and arrangement as in 
U. leucJii. 

External cliaraeiers. — Muzzle averaging shorter and slenderer 
than in Ii. arahievs. IShaj e of ears as in Ii. Jeae^i '■ outer miuh 
more convex than inner margin, tip broad, not attenuated as in 
li. arahinis; aiititragal lohe small, rounded. 

General size neaily always smaller Ihan in Ii. aralicvs : forearm 
80-O-87-5 uim., against 87-90. Eirst digit absolutely shorter: 
length with claw '26-o-2i)-tj mm., against ;-50-;H;i in arubicus ; also 
jiroportioniilly the difference in this respect is well marked : index 
of iiist digit in Ii. lesvlunardti '6'65, in Ii. arahievs li'il. Second 
digit shorter, the tip of its claw in A', hsc/ienaulti as a rule falling 
a little short of, in Ii. arahiens as a rule being on a level with or 
reaching beyond, the tip of the third metacarpal. Metacaipals and 


]ih;iLuip;es of third, fourtli, and fifth dijjifs (except the second 
phiihiiix of the tilth digit) shorter thiui in R. aralncH>i, the dill'ererice 
bein;;; speciailj- notieoat)le in tlie absolute length of the first p'lahuiK 
of the third 'digit (33-8-;3(3'8 mm. in 11. hfchenanlH, 37-39-8 in 
li. (f rahicufi) and the second pliahinx of the third dvjit (41— Kr2 mm. 
in It. Iesch>'nn>dti., oO-[)-(V) in li. (imhicns: index of this phalanx 
in It. I'scheiiaidti 521, in It. arabicns 601). Length of tibia as in 
ii'. nrahicux, but foot averaging smaller : me isured witli claws 
2U-23-5 mm. in li. leschenaulti. 22-7-25-5 in It. arahicus. 

Quality and distribution of fur essenrially as in R. arahinis, but 
tibia more thinly haired. Colour of a darker and browner shade, 
especially on the crown : — Back and rump dark and dull brown 
(darker than Front's brown) ; crown and occiput lirowiiish bistre; 
nape of neck varying from light drab to almost wood-brown; under- 
side between drab and isabella colour. A considerably brighter- 
coloured phase occurs : mars-brown on back and rump, Prout's 
brown on crown, wood-brown on the whole of the underside. 
Measurements. On p. 48. 

Itaiir/e. Himalayas (Nepal), extending southward over the Indian 
Peninsula, eastward through Bengal, Burma, Siam (Laos Mts.) 
to S. China (Araoy). 

Cutijpes in the Paris iluseum. 

rte'ropusleschennulti, Desm.; 1820.— Type locality : " les environs 
do Pondichery." Cotyi)cs, two adult males, mounted, much faded, 
hibolled "Pondichery", Leschenault "' ; Eeg. nos. A. 82 and A. 83; 
skull of no. 82 extracted, of no. 83 in situ. I'laeed by Desmarest 
in the section " Ronssettes sans queue," an error corrected by Is. 
(leoftVoy in 1828, /. s. c, on examination of Ihe type : (" sa queue, 
trcs-visible, u'est qu'a ])eine cngageu dans la membrane inter- 
femorale, et a environ six lignes de long "). 

Pti'rojmx pi/rivorux, Hodgs. ; 1835.— Type locality: the central 
region of Nepal ; cotypes in collection, figured in Hodgson's un- 
pulilislied drawings. 

Cifiiopierns nffini^, Gray ; 18-13. — No description. Two specimens 
were registered by Gray under this name, both from the "Hima- 
laya," viz. no. 130rt [ =], which is the type, still in 
the collection of the British ifuseum, and 130 ft [ =, 
"younger," not found in the collection. Indistinguishable from 
R. leschrnaidti. 

FJcutherurci ftdiip'vosa, Gray; 1870. — Type locality : Laos Mts., 
Siam (Mouhot Coll.); tyjie incoUection.— Brief description of the 
colour of the fur. I am unable to discriminate this specimen from 
It. lesdienaxlti. 

Eh'uJherura fvsca, Gray; 1870. — Type locality uncertain 
("India?"'; purchased from Par/udaki) : tyi>e in collection. — 
Separated by Gray on account of its "much brighter and redder" 
colour. Is the brighter-coloured phase of R. kxchemmlti. 

Cinwnycteris infmeutd. Pet.; Ib73.— Type locality: " angeblieh 
aus Calcutta" (a dealer's specimen); type in the Berlin Museum 
(no. 301 ). " Schr iihnlich der 0. lesrh^naKhii. in alien Verhtiltnissen 



kleiiior, flnnkelbrnnn von Farbe, mit schwarzon Krallcn uud don 
erstoii falscluni liackzalin g-riisser " : detailed measurements given : 
forearm 68, tliiid metacarpal 42, tibia 29 mm. The type, I am 
informed by Trof. Matschie, is a young (not full-grown) individual 
(■'die Epipbysen an den Fingergelenken sind noch nicht mit den 
Phalangen verwachsen," Matschie, in litt.) ; hence its small size. 

Remarks. — li. leschennaUi is at once discriminated from the fore- 
going species by the shortness of the second phalanx of the thi'd 
digit, a character which it shares with all the Eastern species of the 

Piu-cbased (J. Tamer). .38 3.13.37. 
{Ti/pe i)t' Viniojjierus affiim. Gi-aj-.) 
B. if. H.idgsoii. Esq. [P.]. Not reg. 
(Coiypes of Pteriiqiuf, pyrivorus, Hodgs.) 

B.fl. Hodgson, Esq. [P.].<5. 
Cajit. \V. J. E. Boys 4.^.2.1.14. 


Capt. W. J. E. Boys r 
|C.]. [49.8.16.y. 

(e &/ labelled Elcafhcrnra marqiiuifa. Gray.) 

Lieut. E. Y. VVatsun [P\J. 

Si;im. Moubot Coll. 62.8.18.r). 

{Type of F.lcnfhcrvrn fid'iqhui»a . (Trnv ) 

'Pufcliasfd (Pai-ziidaki)! 4!>.8.23.1(). 
{Type of Eleufhrrura j'uFca, Gray.) 

Lidth de Jeiide Coll. (■.7'.4. 12 32f5. 

5. Roiisettus seminiidus, Gray. 

Oynoni/cferis am])le.vkandnta (pt.), Dobson, Gat. 
Chir. B. M. p. 72. 

Pteropus leschenaulti (nee Desni.), Blyili, J. A. S. B. xx. no. 211', 

p. l.')5 (1851 : Ceylon) ; id, op. cit. x.xi. no. 228, p. 34o (1852: 

Ceylon); Kelaart, Prudr. Faun. Zeylan. p. 27 (1852: Ce.ylou); 

Wayner, Schrebrr's Siiuy., Si/ppl. v. p. 604 (pt.) (1853-55 : Ceylon) ; 

Giehel, Siiuy. p. 999 (pt.) (1855: Ceylon); Blytli, Cat. Muvim. 

Mun. A.'t. 8oc. p. 21, no. 54 (}it.) (1863: Cevlouj ; Sterndale, 

Mamm. ImUa, p. 40 (pt.) (1884: Ceylon). 
Xauthaipyia leschenaulti, Fitzinyer, SB. Alcad. Wien, xlii. Heft 25, 

p. 389 "(1861: Ceylon) ; id ^' op. cit. Ix. Abtb. i. Heft 8, p. 472 

(pt.) (1869: Ceylon). 
Cynonycteris (Pleiopus) leschenaulti, Zcklor, Beise ' Koiara,^ 

'Siiiiy. p. 12 (1869 : Ceylon). 
C'ynoiiycteris amplexicaudata (pt., nee Geoff", j, Pefer.f, MB. Akad. 

Berlin, 1867, p. 865 (Cevlon); Ihilmjit, J. A. S. B. xlii. pt. ii. 

p. 202 (1873) ; id., Cat. i'hir.Ind. Mus. p. 2 (pt.) (1874 : Ceylon) ; 

id., Mvn. Asint. Chir. pp. 29, 190 (187f>: Cf-ylon) ; J. Anderson, 

Cut. Ahniim. Ind. Mus. i. p. 103 (1881: Cevlon); Truuessart. 

Cat. Mcnmn. i. p. 84 (1897). 
Cynopterus (Cyuonvcteris) amplexicaudatus, Trouessdrf, Bee. ^• 

May. Zoul. (3) vi.'p. 20(j (pt.) (1878: Ceylon). 


(S fid. sk. ; 



2 d ad. al. ; 
skull ot h. 



Ad. skull. 



2 ad. sk. ; 





2 ad. sk.; 





2 ad., 2 


iiniii. al.; 


.skuU of 'f. 

i.; skull. 



luiui. sk. ; 


2 ad. al. 


Xaiith;ivpyia aniplfxicandata (pt.), Blanford, Fami. Brit. Lid., 

Mamm. pt. ii. p. 201 (18;)1 : Cevlouj ; Matschie, Megachiroplrra, 

p. (37 (pt.) (1899). 
Xantliarpyia tiemiiuida, Gray, Cat. Monk. Sfc. p. 115 (1870: 

Rousettus seniinudus, K. Andersen, A7in. ^- May. N. H. (7) xix. 

p. 538 (1907,1. 

Diagnosis. — Similar to R. leschenaulfi, but p' deciduous, iiajie 
and slioulders scmi-uakt'd, general colour of fur lij^hter. Forearm 
79-8o-5 mm. 

Detaih. — Size and shape of skull as in 7^. JescJienaulii. p' de- 
ciduous* ; m, quite or almost equalling m, iu length ; m^ elliptical 
in outline. Palate-ridges as in Ji. hschenaulti. 

Pur markedly shorter and more closely adpressed than in 
7^. hucheiumltl ; nape and shoulders covered with such sjiarse and 
short hairs as to appear semi-naked ; hairs on membranes and 
upperside of forearm and tibia shorter and sparser : in some 
individuals these parts appear to be almost naked. Colour lighter 
than ill 7i. Icsclienaultl : upperside intermediate between mars- 
brown and wood-brown, or very nearly wood-brown ; head darker; 
underside wood-brown. 

M.-asiiremenis. On p. 48. 

Range. Ceylon. 

Type in collection. 

Xaiitharpijia .'leminuda. Gray ; 1S70. — Typo locality : Ceylon.— 
The name Pteropus semimidits is commonly assigned to Kelaart, but 
Kelaart seems never to have published any description of the species ; 
in the paper usually referred to by authors, viz. Blyth's account in 
J. A. S. li. xxi. p. 3-15 (1852) on a collection of mammals sent by 
Kelaart to the Asiatic Society of Bengal, the name ajipears only as 
a synonym, without comment, of Pt. lescheiianlti, and the same is 
the case in Kelaart's ' Prodromus Fauna Zeylanicae ' (1852). It 
remained a nomen midiirn, until iu 1870 (Z. s. c.) Gray published 
a brief description (quality of fur, colour, length of forearm) of 
" XaatJuirpyia seminada" and the British iluseum specimen on 
which he based this descriiition is, therefore, the type of the 

o. (^ ; skull. Ceylou (l/nvaites). Mr. Cuming's Coll. Not reg. 

(Ti/pe of fpocies.) 
6. cJ ;skull. Ceylon. Dr. Kelaart [C. & P.J. .'i2.5.9.10. 

(Authentic specimen of"' Pteropus semiinuliis. Ke\"^ 
c. J ad. sli.; skull. Punduloya. Cevlou, E. E. Green, Esq. [P.]. 9,j. 7.27.1. 

4:joo' ; 30 Apr. is;i.^. 

* Details from three skulls : — one im'nature, p^ present on both sides; one 
ad., tei'th very slightly worn, p' present on one, nbsent on the other side: one 
ad., tfetli unworn, p' ili>i»nt on both i-'Hoa 


0. Rousettus amplexicaudatus, E. Geoff. 

Cijnonijcteris atiij_)lv.iicaadata (pt.)i Dobsou, Cat. 
Chir. \^. M. J). 12. 

rteropus amplexicaudatus, i'. Geoffroi/, Ann. Mus. d'Hist. Nat. 
XV. p. yC), pi. iv. (whole tif,r) (1810: Tiinoi-j ; Okeii, I.elirb. 
XatunjeKch iii. Abth. ii. p. 934 (1816 : Timor) ; G. Vucier, Ithjue 
Anim. i. p. 1:24 (1617) ; Desinarest, Now. Did. ci' Hi^t. Nut. 
xxix. p. r)13 (1<"?19 : Timor) ; id., JEmycl. Meth., iVJatinit. i. 
p. Ill, no. 145 (1820: Timor); Temmiiwk, Moii. Maiiiin. i. 
p. 200 (pt.), pi. .\iii. (whole iig.), pi. xv. lig. 16 (bead) (182".: 
i'imor, WumaU-a) ; Lei^son, Muii. J^iamm. p. 112, do. 203 (1827 : 
Timor) ; Gir/t/, in Griffith's Anim. Kiti(;d. v. p. 57, uo. 162 
( 1827 : Timor) ; Desmaresf, Diet. Sci. Nut. xlvi. p. 367 (pt.) (1827: 
Timor, Sumatra); Is. Geo/f-oy, Diet. Class, dllist. Nut. xiv. 
p. 703 (pt.) (1828: Timor, iSumatra) ; J. B. Ftsclier, Syn. Mnmni. 
pp. 86, 549 (1829: Timoi', Sumatra); li. Geofroy, Mist. ^ at. 
Mamvi. lei;. 13, p. 25 (1829); Wayler, Syst. d. Aniphihiiii,-^. 9 
(1830) ; Lesson, Hist. N(d. Manwi. (Co7npl. Biiffhn) v. p. 55 (i:t.) 
(1836 : Timor, Sumatra) ; Temvdnck, Man. Mamnt. ii. p. 90, 
pi. XXX vi. H>^s. 18, 19 (sivuU) (1837) ; S. Mi'dler, in 7'evmiinrk's 
Nat. Gesch. Nederl. overz. Lez., Zoof/d. ]ip. 20, 58, 59 (pt.) (1839-44 : 
Timor, Sumatra) ; Waijncr, Sdirebers Siiui/., SuppL i. p. 359 
(pt.) (1840; Timor, Sumatra); Lesson, N. Tail. Ithjue An. 
p. 14, no. li)3 (pt.) (1842: Timor, Sumatra) ; 8ulanz, iSyst. J'erz. 
iSduy. i. p. 131 (pt.) (1844 : Timor, Sumatra ) ; U'ayni-r, Schrehers 
Siiuy., fSuppl. V. p. 604 (pt.) (1853-55: Timor, Sumatra]; Gielxd, 
Siiuy. p. 1000 (pt.) (1855: Timor, Sumatra) ; tSihleyd, Dierhttnde, 
i. p. b'-i (1857: Sumatra); Fatsih, J^eu-Guinca, p. 150 (pt.j 
(1865: Timor). 

PacliYSoma amplexicaudatum, Wntcrhouse, P. Z. S. 1843, p. 67 

Pteropus ((!\niinycteris) amplexicaudatus, Peters, MB. Akad. Berlin, 
18()1, p. 707 (Luzon, Samar). 

Xantbarpvia aniplexicaudata, Gerrard, Ccd Bones Mamm. B. M. 
p. 58 (1862); Fitzuu/er, SB. Akad. Wien, 1.x. Abih. i. Heft 8, 
p. 470 (pt.) (1869: Timor, Sumatra); Thovms, Ann. Mus. Civ. 
(Jenova, (2) .xiv. p. 108 (1894: Eiigano) ; id., Nov. Zool. iv. 
p. 263 (1897: Savu I.) ; id., Trans. Zool. Soc. xiv. pt. vi. p. 383 
( 1898 : Luzon) ; Hurtert, Nov. Zuul. v. p. 456 (1898 : Alor I.) ; 
Matschie, Meyaehiroptera, p. 67 (pt.) (18i.'9). 

Cynt)uycteris ample.xiraudata, Peters, MB. Akad. Berlin, 1867, 
"p. 865 (pt.) (Timor, Philippines) ; Dobson, J. A. S. B. xlii. pt. ii. 
p. 202 (pt.) (1873) ; Peters, MB. Akad. Berlin, 1873, p. 485 (type 
of species re-examined) ; Dobson, Mon. As. Chir. pp. 29, 190 
(pt.) (1876 : rhiiippine;;) ; id., P. Z. ,S. 1878, p. 877 (Cambodja) ; 
id., Pep. Brit. Assoc. 1880, p. 1 73 (Cambodja) ; Rubin, C. R. Acad. 
8ci. xc. p. 1369 (1880: anatomy); id., Ann. Hci. Nat., Zool. 
(6) xii. art. 2, pp. 4lS: seq., pi. v. lig. 30, vii. bg. 47 (organs ot repro- 
duction) (ls81); J.Anderson, Cat. Mamm. Ltd. Mus. i. p. 103 
(pt.) (1881: Pbilippiaes) ; Jeidink, Cat. Osleol. Mamm. p. 263 
(pt.) (1887 : Timor, Sumatra) ; id.. Cat. Syst. Mamm. p. 150 (pt.) 
(1886: Timor) ; D. G. Ediot, Field Coliunb. Mus. Piibl., Zool. i. 
no. 3, p. 79 (1896: Negios L) ; 'J'ronessart, Cat. Maiinn. i. 
p. 84 (pt.j (1897); tieabra, J. Sci. Math. Lisbon, (2) v. no. 19, 
pp. 161, 168, pi. i. lig. 10 (palaty-ridgc.-) (1S'.»8: Timor). 

RorsF.rrns ,\Mri,r,xiCArpATrs. 


('viinii\ct('i'is (I'ternpiis) nniplc.xicauilatiis, Mnvchi, Atti Sic. Ital. 
'■Sii. Ndt. XV. p. 517 (187i'-7.'< : structure nt' Imirs). 

(.'viiopierus (Cynonvcteris) iuii])l(;xicaudatus, Trouessart, licv. iS- Mm/. 
' Zuol. (;J) vi.'p. :266 (pt.) (1878). 

Kousettus fiinplexifaudatu.-;, Troiwsxiivt, Cat. Mamm., Siip/il. p. (iO 
(pt.) (llKU); Miller, Pruc. U.S. Nat. yiiis. xxx. p. 824 (1906: 
Euyaiio) ; K. Anih-rsvn, Ajvi. ^- Ma;/. X. H. (7) xix. p. '"-08 
(1907) ; Miller, Fan,, if Gen. Bats. p. o4 (pt.) (1907). 

Elontlierura int'uniata, Gray, Cat. Monk. ^-c. p. 118 (1870: Flores). 

Eleutherura philippiuen.-is, Graij, Cat. Monk. SfC. p. 119 (1870: 

PuTopus ])lulippineasi.s, Elern, Oit. Sid. Faiiua Filij)inas, i. ]). 6 
(1895: Luzon; Samar ; Min^'anao ; Palawan). 

Itousettus philippinensis, Miller, Fam. ^- Gm. liat^, p. 54 * (1907). 

Cvnoiivcteri.s bocaiiei. Seabra, J. Sci. Math. Lisboa, {'2) v. no. 19, 
'pp. it"0, 1G9, pi. i. tifr. 11 (paktu-ridge*) (1>^98: Timor). 

liouseltus bocagei, Truutssart, Cut. Mumm., Siippl. p. UO (1904). 

IHa;f)Wsis. — Similar to li. leschenauM, bttt m^ suljcireular in 
outline, ears narrower. Forearm 77-S7"2 mm. 

Skull and teeth. — Skull essentially as in Jt. hf'clunaulti, but 
avera^^ing smaller; rostrum proportionally .slenderer. — Teeth 
smaller: maxillary row 12-8-14:2 mm., against 14-15-7 in 7i. /e- 
srJwnaulti. m^ subcircnlar in outline, not elliptical as in It. leschen- 
aitlti. In li. amj^) there is a distinct tendency to a 
reduction of the interspace between the upper canine and p\ i^ 
being in some individuals rather closely wedged in between those 
two teeth ; but the character is not iixed, many examples (inde- 
pendent of geographical habitat) having the interspace c-p' as 
broad, and p' as distinctly separated from either of these teeth, 
as li. lexelienaulti; li. amplexicaudntns forms in this respect a 
transition between R. ItsehenavUi and by-ac/njoiis, in which latter p^ 
is deciduous, c and p'' occasimially in contact. 

r(date-rid</es.—4 + '3+\, as in E. le^rhenaulti, rarely 4 + 4-4-1 
(specimen " b " in the list below). 

Frternal chnracters. — Muzzle averaging shorter and slenderer 
than in li. le.s-rheiianlti: length from tVont of eye to tip of nostrils 
i;$-5-15-5 mm., against 14-8-16 in li. lescJienanlti. Ears much 
narrower: greatest width 12-13, ag-ainst 14-5-15-5 in li. Icschen- 
aidti; tip of ear not attenuated; antitragal lobe small, rounded. 
Tail averaging longer, tibia markedly shorter than in li. lefchenaidti. 

Quality and distribution of fur as in It. UscJienaalti ; upperaide of 
tibia almost naked. Colour of fur as a rule distinctly darker than 
in li. lesrhinatdti: head, bark, and rump varying from dark Trout's 
brown to dark olive or sepia-brown ; nape from wood-brown to 
broccoli-brown ; sides of neck and forencck more or less suffused 
■with wood-brown or lawny olive in males, broccoli-brown in females ; 
chest, breast, and belly dark greyish drab, sometimes inclining to 

Measurements. On p. 48. 

* Spelt Rousptlu!' phillppeiifig in the iiuk-x (p. 276). 



Range. Carabodja, Piiiliiipiues, Borneo, Sumatra, Engano, Flores, 
Savu, Alor. Timor. 

Tjipe in the Paris Museum. 

Fterojms am/>Je.cicaiulatn.s, Geoff. ; 1810. — Based bj- Geoffroy on 
"plusieursindividus" obtained in Timor during Peron and Lesueur's 
voyage (Capt. Baudin) ; only one of these appears now tc be in the 
Paris Museum, a young individual, mounted, much faded, skull 
extracted, labelled " Timor, Exp. Baudin" ; lieg. no. A. 79. 

ELeutlierura infumata, Graj' ; 1870. — Type locality : Flores 
(A. 11. Wallace) ; type in collection. — The name would seem to 
indicate that Gray separated the Flores specimen on account of its 
" blackish browu "' colour ; it ditfei's, however, neither in this nor in 
other respects from a majority of examples oi R. amphxicaudatus. 

Eleulherura pJiilippinensis, Gray; 1870. — Type locality: Manila 
(Hugh Cuming); type in collection. — Gray gives a brief description 
of tbe colour of the specimen, without pointing out his reasons for 
separating it from Pleropus amplexicaudatus, Geoff. The type, 
as well as more recently acquired specimens from the Philippines, 
differ m no respect from R. amplexicaudatus. 

Ci/7ioni/cteris hocar/ei, Seabra ; ISUS. — Type locality : Dyli, Timor 
(Fr. Newton); type in the Lisbon Museum. ^ — Separated by Seabra 
from R. ahip>lexicaudatus on account of the supposed greater 
zygomatic width of the skull (and a trivial difference in the form 
and position of the sixth and eighth palate-ridges). Zygomatic 
width of the type skull {S ) not given ; in the figure {I. s. c. pi. i. 
fig. 11) it measures scarcely 23 mm. ; in a British Museum 
specimen ( c? ) from Alor I., north of Timor, 24 ram., in another 
( $ ) from the same island 21*2 : similar variations are found in 
R. amplexkaudatus and allied species from an)' locality. 

Remarlcs. — R. amplexlcaudaius is readily distinguished from 
R. lescJienaulti by its much narrower ears and the difi'erent shape 
of nig. ill. hschenaidti is continental in range, R. amplexicnudatus 
chiefiy Indo-Malayan (insular) ; their areas probably touch each 
other somewhere in S.E. Asia ; R. leschcnaidti is represented in the 
collection from Burma and Siam, R. ample.clcaudatas from Cam- 
bodja. — Some authors (Peters, Anderson and de Winton) have laid 
stress on the greater length of the tail in Indo-Malayan specimens 
{amplexicaudaius) as compared with examples from tlie Himalayas 
and the Indian Peninsula (lescJienaulti) ; the tail averages, in fact, 
decidedly louger in wmplexicaudcitus, but the character, as being 
subject to a good deal of individual variation, is practically not of 
much use for a discrimination of the two species. 

a. Ad. sk. ; skull. Cambodja (iVou- Tomes Coll. 

hot Coll.). 

b. 2 imiii. al.; Philippines. Zool. Soc. Coll. Not reg. 


c. S ad. sk. ; Manila. Luzon. Hugh Cuming, Esq. 

.skull. [C.]. 

(Type oi Elci'.lherura philip2>ineHsu, Gray.) 


d-e. S 'tI" " '«'• E<'iiKuet. N. Lu- J. Whiieliead, Esq. OT.o.^.o-C. 

sks. ; skulls. zm, i^lHJO' ; [C.j. 

14 Febr. 18i)4. 
./-.9. c? iw^.. (5 P"ll- Banuii, ISarauak. Dr. Chas. Koso [P,]. 0.7.29.'2-3. 

al. ; skull 


//. ^ acl. sk. ; Suinalra. Purchased (J. Tiir- 3S.3.13..'ib. 

skull. iier). 

i. 2a(; skull. Bua-Kua, En- Marquis C4. Doria 
gano {Dr. K. [P.J. 

j. skull. Flores. Dr. A. R. Wallace (i3.12.26.12. 

{T(ipc of Eleafhrrurti hifumata, Gray.) 

>!•. ;skul!. Saru I.. W. of A. Everett [C.]. 

Tiincir ; Aug. 

l-m. d' ad., 9 ad. AJor I., ^\ of A. Everett [C.]. 

sks. ; skulls. Timor; Mar. 


n. luiui. St. Tiuior. Purchased (Frank;. 44.4.4.(5. 

7. Rousettus minor, Dohs. 
Ci/iionycteris minor, Dobson, Cut. Cliir. B. M. p. 73. 

Pt-n-opn.s aniplexicaudatus (pt., nee Geoff.), Besmaypst, Diet. Sci. 

Nat. xlvi. p. 367 (1827 : Java) ; Lesson, Hist. Nut. Mumm. {Conipl. 

Hufon) V. p. o5 (18-JG : Java) ; 8. Miiller, in 'i'eiiiniitick's Nut. 

Gesch. Nedeii overz. hcz., Zuoyd. pp. 20, 58 (1839-44: Java); 

Les><on, N. Tall. Rhyne An. p. 14, no. 193 (1842: Java); 

Schinz, Syst. Verz. Siiuy. i. p. 131 (1844: Java); Wayiier, 

Sc/irehers Sihiy., Suppl. t. p. 604 (1853-55: Java). 
L'vnonvctoris atii'ple.\icaudata, Ji^/^^iwA-, ^Teber's Zuol.Eryehn. Niederl. 

' Ost-Iud. Ileit i. p. 12(5 (I8i : Java). 
Xaiitliarpyia ample.xioaudata (pt.), Ji/i/th, Cat. Mamm. 3Im. As. Sac. 

p. 2] , no. 55 (1863 : Java) ; Mutschie, Meyachiroptera, p. 67 

(1899: Java), 
llou.settus aniplexicaudatus (pt.), Trotiessart, Cat. Mumm., Suppl. 

p. 00(1904: Java). 
C^nonvctevi.t luinor, Dobson, J. A. S. B. slii. pt. ii. p. 203, pi. xiv. 

fi^. 9 ( (1873 : Java) ; id.. Cat. Chir. Ind. Mus. p. 2 (1874 : 

" Malay Countries ") ; id., Mon. Asiat. Chir. pp. 32, 190 

(187U: Java); J. Arulerson, Cut. Mamm. Ivd. 31us. i. p. 104 

(1881 : Java) ; Trouessart, Cat. Mamm. i. p. 84 (1897 : Java). 
Cvnoptents (Cyuonvcleri.s) niiuor, Trouessart, Jieo. Sf May. Zool. 

■(3) vi. p. 20(5 (1878 : Java). 
Rou.«i'ttu.s minor, A'. Andersen, Ann. ^- Ma//. N. H. (7) xii. p. 509 


The only specimen I liave seen is the type in the Calcutta 
Museum, a' dried skin (perloctly adult, tectli almost un\yorn), in 
bad state of preservation ; colour faded to pale rus.set ; basioccipital 
and sphenoid portion of skull wantinj,'. The precise characters of 
the species cannot be given from this material. 

SLitll and tetth.— f^knll not differing appreciably from that of 
11. brachifotis, except perhai)s in the .slightly larger size of the orbits 


(only one (|nite fiill-sizcfl skull of brachi/otis available for com- 
parison). oiil}' tangible difference in the teeth would seem to 
he the slightly smaller width of the premolars and molars in 
li. minor, but the character must be taken with caution, inasmuch 
as there is some individual variation in this respect in 11. hrachi/vtis. 
p^ a little smaller in bulk than an inner upper incisor, separated by 
miuute spaces from the canine and p'^. ni^ subcircular in outline 
(a little longer than broad). 

Rvternal characters. — According to Dobson the ears in R. minor 
are much longer than in li. bracJu/oiis (see his description of 
li. hrac7i(/otis, Cat. Chir. p. 74). Tliis statement is erroneous ; the 
oars in the type of i?. yninor (the left ear-conch is well preserved, the 
riglit much damaged) are of the same length as in dried skins of 
li. hrachi/otis, or if there is any difl'erence, they are perhaps a trifle 
shorter ; the width of the ears cannot be estimated with certainty, 
but would seem to be slightly smaller than in hrachifotis. In the 
relative length of the muzzle (wrongly stated by Dobson to be 
shorter than in brachi/oiis), the distribution of the fur, and the 
measurements of the forearm, metacarpals, phalanges, and tibia, 
there is no difference from li. braclnjotis. 

Measurements. On i). 4S. 

liange. Java. 

Tjiiie in the Calcutta iluseura. 

liemarks. — Judging from the single specimen examined, li. minor 
appears to be more closely allied to It. brachi/oiis than to li. amplcvi- 
caiulattis. From the latter species it is readily distinguislied by 
its smaller size and relatively smaller ears. Its differences from 
M. brachyotis remain to be determined *. 

8. Rousettus brachyotis, Bobs. 

Cjinonycteris brcicJiyotis, Dobson, Cat, Chir. B. M. p. 74. 

Pt'.^ropus amplexicaudatus (pt., nee Geoff.), Desmaresf, Diet. Set. Xat. 
xlvi. p. yi)7 (18:^7: Ambnina) ; Is. Gfofroif, Diet. Class. d'Hisf. 
Nat. xiv. p. lO'.'j {\S'26 : Amboina) ; J. B. Fiseher, Sj/n. Manim. 
p. SO (1829: Amboina); Lesson, Hist. Nat. Mamm. (Compl. 
liiiffiit)) v. p. oo (183(i: Amboina); <S'. Miiller, in Temminck's 
Naf. Gesch. Nederl. overz. hez., Zoogd. pp. 20, 59 (1839-44: 
Ani))oin,i) ; Ww/ner, Schrehers Siiin/., Suppl. i. p. 359 (1840: 
Amboina); Lesson, N. Tahl. Rec/ne An. p. 14, no. 193 (184:^: 
Amboina); Sc/iinz, Si/st. Verz. Sauij. i. y. 131 (1844: Amboina); 
Wagner, Sehrebe/s Sitiig., Suppl. v. p. 004 (]853-5o: Amboina) ; 
Giebel, Sdug. p. 1000 (1855 : Amboina) ; Fiiiseh, ISeu-Guinea, 
•■ Vj p. 150 (1865: Amboina). 

* While these pages wfire going through the press, a fine series of skins 
and alcoliolic specimens of Rouselius miiwr obtained in Java by Mr. Giij 
C. Sliortridge were presented to the Museum br Mr. W. E. Enlston. Full 
description of the species based on lliis mattriul will be given in the ' Addenda ' 
to tliis volume. 


Cviion7ctfvi.« ani])loxicaii(la(ii {i\t.), I'cfcrs, Mil. Al-ad. />,r!,'n.]fi(u, 

]i. si).") fAiuboina); Jentink, Cut. Si/st. Maiiun. p. 1")1 (]8r!8: 

Xaiilliiiri)\ia amplexicaiidata (])t.), T'ltziiK/er, SB. Ahod. ]]'ien, Ix. 

Abih. i". lleCr ■■<, p. 470 (1860 : Amb.iiiia). 
CvnMpii,'ius (1 viioiivcieri.s) airplexicaudatus, Troiiessdrt, Iin\ S; 

' M<i(/. Zi.ul. (?>) vi. p. l'OG(pt.) (1878: Aiuboina). 
Cvnonvctm-is brachvcti^, I)ofj.<on, P. Z. S. 1877, y. IKi (Duke of 
'Yoiiv I.); id.,' P. Z. S. 1878, p. 816 (l)uku of Voik I.); 

Th'imns, P. Z. S. 1887, pp. 3i'3, .'5l'7 (Solnmou Is.) ; id., I'. Z. 8. 

1>^f^t<, (ip. 47o, 483,484 (.Soh.iiiun Is.) ; Jcntiiil-, Cat. S'/ft Mcotim. 

p. h")! (1888: Duke of York I.); 'J ruuessart, tat. Mainiu. i. 

p. 84 (pt.) (1^97). 
C}noptorii.'3 (Cviiouvcteri.s) braclivotis, Truues-sart, liev. 4'- 3/"'/. 

Zool. (3) vi. p. 201') (1878). * 

Xanthai'pyia brachvotis, Mat.'uhie, Mcf/achiropterti, p.fiS (pt.) (1899 : 

New Guinea, New Ireland, .Solomon Is.). 
Ilousettus braclivotis, Trouessart, Cat. Mamii)., StippL p. 60 (]it.) 

(1904) ; K. Andersen, Ann. if Mar/. X. H. (7) xix. p. .3UD (1907) ; 

Milk-r, Fam. ^- Gen. Bid>:, p. o4 (1C07). 

Diiicjiio^is. — ISiiuilar to JL ampli'dicaxdatus, but smaller, with 
shorter and narrower ears ; p' deciduous ; toutli-rows shorter. 
Forearm 70-75 uim. 

k</,ull and teeth.— SliviU averaging smaller tliau in II. amjdt\ci- 
cniidaius, and with noticeably slenderer rostrum, but otherwise not 
difl'ering in shape. — p' deciduous * ; it' jiresent, as a rule elostly 
wedged in between the canine and p'^ ; interspace c-p^ as a rule 
narrower than in any of tlie foregoing species, the two teeth being 
occasionally in contact. Cheek-teeth more crowded ; p'' and molars 
proportionally quite as heavy as (if not heavier than) in li. aiitjjh'.ri- 
candatus; m., maikedly shorter than m^ ; m^ subcircular in outline, 
as ill 2i. ample.vicaudatus. 

Palate-rii/(/i's. — ^ + 4+1; one specimen only examined; some 
slight individual variation may occur. 

E.vternal characters. — General si/e smaller (at least on an average) 
t\\&n\n It. amj^ilexitaudatus \ see table of measurements. Ears .short 
and narrow, but scarcely ditiering in form from those of R. 
caudatus; not attenuated below the tip, the tip itself broadly 
rounded off; antirragal lobe small, rounded. 

Quality and distribution of fur as in 11. ample.vicaudatus ; tibife 
practically naked above ; jiro.ximal portion of forearm thinly covered 
with short hairs. Coloration as in the darkest specimens of 
Jl. aiujilc.vicandatus. 

Meusurouents. On p. 48. 

Itamje. Amboina ^Genoa Mus.), New Guinea (Berlin Mus.), 
Bisnuirek Archipdago, fSolomon Islands. 

Tiii>t in collection. 

Remarks. — This, the most eastern representative of the genus, is 

* Details from six skulls: — one jiiT. (not quite full-grown), two youug 
adults: p' pre.-ent on both sides; one young adult : p' pre.seut ou one side 
CiAy ; two adults (teeth slightly worn) : p' absent on botli sides. 


very closely allicil to lu ampJe.vlccmdaUis, dilleriiig only in its 
smaller size, i)roportionally smaller ears, slenderer rostrum, de- 
ciduous p\ and (as a rule) narrower interspace between c and p'. 
T!ie western limits of its range (islands VV. and S.W. of Nevv 
Guinea) remain to be detei-mined ; a specimen in the Genoa 
Museum from Amboina (collected by 0. Beccari, and examined by 
the present w liter) is E. hrac/ii/otis, but its alleged occurrence in 
Celebes (Jentink, Notes Leyden Mus. v. ]>. ITo, l^So) probably 
rests on confusion with a distinct species {JL cfle/>e)i.sis, infra). 

a. 2 ^^- sl^-' Unlie of York I. ]{,ev. J. Brown 

skuU. !<.'.]. ( 7//;?P of species.) 

b.; Duke of York I. Ee.v. J. Brown 77.7. 1«.4. 

skull. [C.]. 

c. cJad. sk-; Duke of York I. Rev. J. Brown 7.S.15.5. 

skull. [C.]. 

d-e 2jun. sks. ■ Fauro, Solomons ; 0. M. Woodfortl, S7. 1.18.4-5. 

'skulls. Mmv 18S(;. Esq. [C.]. 

/. Jun. sk. Gn;ul'i,,!can:u-, J. Meek, Esq. [0.]. 1.11..5.4. 
Solomons ; 12 May, 

9. B,ousettiis celebensis, A'. An J. 

CiinoiiUctcris ample.ricnuihitd, (pt. ), Dobsou, Gat. 
Chir. B. M. p. 73. 

Cyiionycteris amplexicaudata (pt., nee Geof.}, Dohson, J. A. S. B. 

xlii. pt. ii. p. 20:^ (187-5 : Celebes); Jentink, Cat. Osteul. Mamm. 

p. 2C>S (1887: Celebes); id., Vat. >Si/d. Mamm. p. 150(1888:, 

Celebes); Trim/'snart, Cat . Mumvu i. \). S-i (1697 : Celebes). 
Roiisettus amplexieaudatus (pt.), Trouesmrt, Cat. Mamm.. .Snpjil. 

p. GO (1904: Celebes). 
Cvuoptenis (Cvnonvcteris) amplexieaudatus (pt.). Truuef<mrt, Jiur. 

">S- Mm/. y<wL(3} vi. p. 206 (1878: Celebes). 
Cyiionycteris bracbyotis (nee Dohs.), Jentink, Note--> Leyden 3f)i.'<. v. 

■p. 17;j (1883 : N. Celebes) ; Trouessart, Cat. Manaa. i. p. 84 (pt.) 

(1897: Celebes). 
Xantbarpyia bracbyoti.s (pt.), Mat.schie, Mer/ackiroptera, p. 68 (1899: 

Kousettns bracbvotis (pt.), Troncsfiaii, Cat. Mamm , Snppl. p. 60' 

(1904: Celebe.^). 
Cvnonvcteris minor {nac Dobs.), Hickson, A Naturalist in A'. Cclchcs, 

'p. 84 (1889: I.). 
Xantbarpyin minor, A. B. Meyer, Abli. 3Lis. Dresden, vii. no. 7, 

p. ^(iNgO: X. Celebes, iSangbir). 
R-ousettus celebensis, K. Andersen, Ann. Sf Mag. iV. IL. (7) xix. 

p. 509 (June 1, 1907: Mt. jMasarang, Celebis). 

Diagnosis. — Bony palate narrower than in any of the foregoing 
si)ecies; last jjremolar and molars, above and below, unusually 
narrow. Eur longer and richer ; notopatagium partly (or wholly) 
hairy ; general size small. Forearm 72-5-7 >'> nim. 

,'^l-ull. — General size as in Ii. ani2ih.ricau</alas; rostrum very 
low and slender; premaxillaries in simple contact in front ; bonv 


])alate unusually narrow jjosteriorly : width extornally across 
m--m'"' !)•? mm. (two adults), against 10-2-1 1-8 in aniple.vicKvdntus 
(ten adults) ; frontal region between postorbital processes flat ; 
temporal fossa narrow ; temjjoral ridges forming a lov/ Sigittal 
crest in fully mature individuals. 

Teelh. — Upper canine and p'' widely separated ; p' in the centre 
of the inters|)ace between these two teeth, not deciduous ; last 
premolar and molars above and Vielow very nanow : m^ at least 
twice as long as broad ; m^ small, less than half the size of m' ; p^ 
three or four times the size of a lower incisor; m.^ about half the 
length (or less) of m^ ; m, subcircular in outline. 

Pdlatc-riihjes. — 4 + '^+ 1 ; arrangement as in li. cunple.vlcaudafus. 

Ears, u'bu/s, tail. — Ears essentially as in li. (implea-icdudaUis: 
narrow, not attenuated below the tip, "the tip itself broadly rounded 
oft' ; antitragal lobe small, rounded. General size of the animal as 
ill li. bracJiifotis (smaller than E. ampIe.ricaudtUus), but digits pro- 
portionally longer than in any other eastern species of the genus 
(index of poUex 392, of third digit 1(546, against 33-5-41 and 
1529-41 respectively in all other eastern species); for details see 
wing-indices, above p. 20. Tail long, probably about 20 mm. 
(only dried skins examined). 

/'MJ-.— Longer, richer, and more velvet than in 11. ctiuidexicaudatus 
and allied eastern species ; notopatagium clothed with dense fur : 
hairing on forearni.s, tibia^, interfemural, and underside of lateral 
membrane longer and richer ; face more densely haired. 

Colour. — Brighter than in B. ampJexicandatus. Back light 
Trout's brown, rump more inclining to mars-brown tinged with 
russet ; sides of back and tibial next to membranes almost vandyck- 
hrown ; crown and occiput dark brown, approaching bistre ; nape 
of neck broccoli-brown ; a glandular tuft of mummy-brown hairs on 
each side of the neck in both sexes ; entire underside of body dark 
greyish drab. — Immature individuals are similar in colour to adults, 
but without the mummy-l)rowu neck-tuft. 

Measurements. See table, p. 48. 

llatif/e. Celebes; Sanghir Islands (Dresden Mus.). 

Tt/2^e in collection. 

liemarlcs. — Without close inspection this species, owing to its 
small size, may be easily (and has in fact repeatedly been) confused 
with It. brac]ii/otis. The larger skull, very narrow palate, narrow 
molars, non-deciduous p\ much longer poUex (28-30 mm., against 
24-26 in hracJi;/otu-), longer wings (chiefly owing to the longer meta- 
carpals), much louger fur, hairy notopatagium, and much more 
densely haired tibia readily distinguish it from B. brachi/ctls, 

rt. c? ad. sk. ; skull. Rurukan, N. Celebes ; Dr. Clias. Hose [C.]. 97.1.2.t>. 

Oct. 185)5. 

A. linni. sk. : skull. Jit. Masaraiig, N.Celebes, Dr. Chas. Hose [C.]. 

4<)W)'; Oct. IWt.i. 

<;. 5 ad. sk. ; skull. Ml. Ma^arang, N. Celebes, Dr. Clias. Hose [C.]. 97. 1 .2.8. 

3.300' ; Oct. 1895. ( Ti/pe of species.; 




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10. Ronsettiis (Stenonycteris) lanosus, Thos. 

Xantharpyia ffigyptiaca (wee Geof.), Horxfield, Cat. Mamm. Mus. E. 

hid. Co. p. 29 (iSol : Abyssinia). 
Eleutherura ae-ryptiuca (pt.), Gruy, Cat. Monk. Sfc. p. 117 (1870: 

Cynonvcteris a?gvptiaca, Dohsim, Cat. Chir. B. M. p. 7o, specimen e 
^ (1878: Abyssinia). 
Cynopterus (Cynonyrteris) ajsryptiaca (pt.), TroufSiart, Rev. i)- May. 

Znol. (.3) vi. p. 206 (1878: Abyssinia). 
lioiisettus peo-yptiacus (pt.), Trouessart, Cat. Mainin., Sappl. p. GO 

(190i: Abyssinia). 
Eousettus lanosus, Thomas, Ann. ^- 3Iag. N. H. (7) xviii. p. 137 

(Aug-. 1, 1906: Ruweuzori East): K. Anderson, n/j. cit. (7) xix. 

p. oil (1907). 

Diagnosis. — Basicrauial axis strongly deflected ; molars exees- 
sively narrow. Antitragal lobe obsolete: wings from back of 
second toe; second phalanx of litth digit longer than (irst phalanx ; 
fur long and coarse; uotopatagium hairy. Forearm 88-5-90 mm. 


Fig. 3.— Rouset/ Its {Steiwni/cicrii;) lanosus, (^. Bmvenzori East. No G 7 1 •■> 
Type. 1. 

SkitU (fig. 3). — General size as in Jl. hachi and anc/o7ensis, but 
more delicately built, the bones thinner and lighter. Brain-case 
80 strongly deflected that the alveolar margin projected backward 
passes through middle or upper half of supraoccipital. Eostrum 

."(I RorsETTrs LAXOsrS. 

sloTulor ; ]vrenia\illaiics in simple contact (uot co-ossiiiecl ), and 
produced distinctly more forward than in li. Itcichi; incisive 
foramina, therefore, longer and broader than nsual in the genus. 
Owing to the strong reduction in the size of the cheek-teeth, the 
temporal muscle is weaker, the teraporal fossa markedly shorter 
and narrower than in B. Jeachi and aii</oJensis, the rami of the 
mandible slenderer, the coronoid process somewhat lower, much 
slenderer, and directed considerably more obliquely backward, 
the condyles smaller, the angular process less projecting. 

Ti-elh "(fig. 3). — Chief character : the extremely narrow cheek- 
teeth. Upper incisors smaller than in li. leuchi and mvjolensis; 
i'-i' rather more separated than i'-i' ; owing to the projection of 
the premaxillaries, the distance i'-c is as great as between c and ]/'. 
p^ as small as an upper incisor, situated nearer to the canine than 
to p''. p''-m^ and p,-!!!^ lower, shorter, and very much narrower 
than in 11. leachi and angoUnsis: width of p'* only about one fifth 
(in all other species about one third) the distance between inner 
sides of p''-p''. m' equalling or slightly exceeding p'' in length, 
m'- about one third the bulk of m'. i^ in cress-section distinctly 
larger than i,. p^ twice or three times the bulk of an outer lower 
incisor. m„ comparatively smaller than in R. leachi and auqoUnsis : 
length about half that of m^. m^ elliptical in outline, about two 
thirds of m,. 

Falate-ridyes. — Examined in one specimen only: 3-f-4-f 1 ; 
arrangement as usual (see p. 19), except that the 5th ridge termi- 
nates closely behind m* (not at m" or between m' and m"), and the 
()th ridge is incomplete, not reaching the lateral margins of the 

/frt^-s^—Similar in length to those of 7?. leachi, but rather 
narrower; outer margin slightly, but distinctly, concave below tip, 
the tip itself narrow, rounded. Antitragal lobe obsolete. 

Wings. — Inserted on base of second toe. 9-10 long vertical 
fasciae in the lateral membrane, viz. 2 postanconeal, 7-8 preancoiieal 
(two alcoholic specimens examined). Wing-srructure essentially 
as in li. leachi, except that the poUex is longer (index 400, against 
361 in /f«c7/i),and the second phalanx of the tiftli digit considerably 
longer than tlie first phalanx. See wing-indices, supra, p. 20. 

Far. — Face short-haired. Fur on body unusually long and 
coarse. Above, proximal two thirds of forearm densely liaired ; 
notopatagium and plagiopatagium next to body, the whole of the 
tibia and the interfemoral, a narrow inner line excepted, furred, 
the hair being particularly long and coarse on the proximal half 
of the tibia and the interfemoral ; upperside of feet covered with 
short hairs. Below, proximal two thirds of forearm, plagiopatagium 
next to body, tibia almost to the ankle-joint, and interfemoral, a 
narrow inner border excepted, clothed with long woolly hair. 

Colour. — Back and rump dark brown, approaching seal-brown, 
inclining to bistre on the posterior part of the rump, interfemoral, 
aiul tibia; crown rather darker than back ; nape of neck between 
br()Ccoli-browu and hair-brown ; the wliole of tiie underparts dark 
urevisli drab. 

RoUSinTl'S ANGOLliXSlS. 51 

^/lasll)^elllenh■. Uii p. 54. 

J{ant/e. Shoa ; lluwcuzoii East (5UU0-13,000'). 

'J'l/pe in collection. 

Eemar/,-s.—-Thia peculiar species, the type of the subgenus Steiio- 
nj/cteris, is chietly characterized by the thin, almost papery condition 
of the bones of the skull, the unusually strong deflection of the 
brain-case, the extremely narrow cheek-teeth, the practically 
complete obliteration of the aniitragal lobe, the insertion of the 
wings on the second instead of the first toe, and the long and coarse 
t'ur. Though much reduced in size the molars do not difi'er in 
structure from those of otlier species of the genus. In the quality 
of the fur It. laiwsus is unlike any species oi liousettus, but closely 
similar to the coarse-haired species of Pteropus. 

c7. Iiiiiu. sk. ; skull. Slioa. Sir W. Cornwallis 61.2 30 6. 

Harris [C.]. 
h. Jatl. al.; -kuU. Ruwenzori East. Eiivvenzori Ex- 

13,000' (H. B. ploration Coinm. 

Wooinum). [P.]. 

c-U. !;^ail., 5;? jnv. ill. Ruwenzori p:ast, Ruwenzori Ex- 
12..5()0- 13,000' ploration Comm. 

{P. B. Woomiam). fp.]. 

e. $ inini. sk. ; Ruwenzori East, Ruwenzori Ex- 6.124 11 

skull- 5000"; 13 Man -h, ploration Coniuj. 

I'.IOO (R. E. Bent). [P.]. 

11. Rousettus (Lissonycteris) angolensis, Boauje. 

Cynonycteris jegvptiaca, Bocaye, Joni. Sri. Math. Lishoa, (2) i no 1 

p. 15 (1889 : Pmigo Andongo). " ' 

? Cynonycteris unicolor (nee Gray), Matschie, Arch. Xatitra i 3 

p. 351 (1801: Cameroon). ' J- • • 

Cynonycteris sp., Bocar/e, Jorn. Sci. Math. Lhhoa, (2) ii. no. 7, 

p. 174, lig. 2 (palate-ridges) (1892 : Pungo Andongo, Cahata,' 

Xaiitharpyia collaris (nee III.), Matschie, Sam. I). Ost-Afrika,v. 17 

(1895: Tanga, Bukoba). ^ 

Cynonycteris collaris, SJiistedt, Bih. K. Sv. Vet.-Akad. Hamll. x.viii. 

Afd. iv. no. 1, pp. 13, 15 16. 46 (1897 : Cameroon) ; id., Mitth, 

Deutsch. Schutzgeb. x. Heft 1, p. 7(1897: Cameroon). 
Rousettus coUnri.s, Thomas, in U. H. Johnstons ' The Uqanda Pro- 

leclorutc; l.\^.4■22 (\m-2). 
Cynonycteris angolensi.x, Bocaye, Jorn. Sci. Math. Lisbua, (2) \-. 

no. 19, p. 133, 138, text-lig. (palate-iidges) (1898: i'lmoo 

Andongo, Cahato, Quibula) ; Seahra, ibid. pp. 159, 109, pi.' i. 

tig. 9 (palate-ridges) (1898). 
Xautliarpyia (Myouvcteri.s) angolensi.'*, Mats,hi/>, Mcnachiroplera 

p. G4 (1899: Angola, Togo, Jjukoba). 
Rousettus angolensis, Troue.<imrf, Cat. Mamm., Sappl. p. 59 (1 90 J) 

A'. Andersen, Ann. Sf May. X. H. (7) \\x. p. 510 (1907). 

2)(£ff/H0.sts.— Ba.sicranial axis very slightly deflected; frontal 
region between postorbital processes distinctly concave; pre- 
maiillaries co ossihed iu front ; molarlform teeth short and broad 



]>, subequal in bulk to a lower incisor. Antitragal lobe distinct; 
wiugs from back of second toe ; lower leg ver_v short (2^-41 mm.) ; 
lur long and silky ; notopataginm hair}-. Smaller than R. Jeaclii : 
forearm 78-83'5 mm. 

Skull (fig. 4). — Brain-case only very slightly deflected, the 
alveolar line if continued backward passing below, or through, the 
lower margin of the occipital condyle. Occipital and hinder parietal 
region considerably more flattened than in any other species of the 
genus, supraoccipital much lower, outline of lambdoid crest, in back 
view, therefore more flatly convex. Owing to the combined eff"ect of 
the slight deflection of the basicranial .axis and the flattening of the 

Fig. 4. — Bo-iiKetfiis (Lisscnyctcris) angolends, J. Riiwenzori Eas-t. No. 6.1"2.4.ri. 
|. (A supernumerary upper molar present on one side in the skull 
figured has been omitted in the drawing ; cf. antea, p. 18.) 

occiput, this latter portion of the skull is directed more backward, 
less obliquely downward, than in other species. Premaxillaries 
firmly co-ossified in front, forming an unbroken bar without any 
trace of a median suture, even in slightly immature individuals ; 
ascending rami of premaxillaries slenderer than in other species, 
particularly in their up])er half. .Supraorbital margin and bases of 
postorbital processes more raised above the le\cl of the upper surface 
of the skull, making the frontal plateau between the processes very 
distinctly concave from side to side. Palation border and meso- 
ptei'vgoid fossa rather narrower than usual. 

RousnTTUs AXGni.KNsis. 0:3 

Ticlh (tig. 4). — c-p'' aiifl i>''-])', as well us c p, and \>-\\ more 
broadly separated than iii li. Icnchl. p' and pj more reduced in 
size, p, in cross-section only equalling or slightlj' exceeding a 
lower incisor. Molariform teeth, above and below, peculiarly 
short and broad, in the upper jaw almost squarish. 

Pulate-ridi/es. — H-\--i-\-2 (three of four alcoholic specimens) or, 
the fourth ridge being not interrupted in the median line, 4-|-3-|-2 
(one specimen). 

Ears. — Outer margin of ear-conch very slightly attenuated below 
the tip, the tip itself broadly rounded off; anr.itragal lobe well 
developed, triangular, subacute. 

Wiiir/s. — Inserted on back of second toe. 12-15 long vertical 
fascise in the lateral mcmbratie, viz. 3—5 postanconeal, 8-1 1 pre- 
anconeal (five alcoholic specimens examined). Forearm shorter 
than in li. leachi : 7>S-83-5 mm., against 89-99. All digits pro- 
portionally considerably longer than in R. leachi : itidex of pollex 
440 (361 in leac^n) ; second digit (index 786) decidedly longer than 
third metacarpal (index 705 ; in leachi respectively 652 and 644) ; 
second phalanx of fifth digit as a rule longer than first ])halanx. 
For further details see wing-indices, supra, p. 20. 

Tail and hind limb. — Tail shorter than in R. leachi : S'5-13 mm., 
against 15-19. ],ower leg much shorter: 2S^-33 mm., against 
40-42 in R. leachi. 

Far. — Ears naked posteriori)', except at base. Face in front of 
and below eyes much more strongly haired than in R. leachi. 
Fur of body much longer, softer, more silky. Distribution of 
fur on humerus and forearm, above and below, as in R. leachi. 
Notopatagiuni densely haired. Upperside of femur, tibia, and inter- 
femoral (a narrow portion next to caloar excepted) long-haired. 

Colour. — Different from that of li. leachi : general aspect, chest- 
nut above, dark wood-brown below. 

Adult male. — IJack and rump rich brown, approaching chestnut 
or burnt-umber. l)ase of hairs light drab or wood-brown, sometimes 
with a tinge of fawn ; crown dark grizzled brown ; fur on forearrr., 
sides of back next to membranes, interfemoral and tibia usually of 
a brighter tinge, approacliing mars-brown or russet. Breast and 
belly wood-brown ; throat and foreneck dark mars-brown tinged 
with fawn. — The bleached coat ( cj ad., liuwenzori East, ilareh 13, 
no. presents a very different aspect : back and rum]) 
rich tawny olive : crown and occij)ut dark raw umber ; breast and 
belly as above : throat and foreneck cinnamon. 

Adult female. — Similar to the male, but throat and foreneck 
scarcely differing in colour from the i-est of the underparts. 

Mea.'<iire))ieitl.s. On p. 54. 

Rantje. Angola, north-westward to Cameroon and Togo, r-ast- 
ward through the Congo Basin td Uuwenzori, and Oerium Kast 

('oti//)f.s in the Lisbon and British Museums. 

lienuirl-s. — R. ant/olcnsi.^, the type of the subgenus Lissoiu/cteris, 
is the most aberrant species of Roasellns ; in none of its peculiar 
characters is it approached by any other species of the genus. In 



the slight deileeliou of the brain-case and the shape of the occipital 
and hinder parietal region it shows distinct leanings towards the 
genus Epomophorus. 

a. 2 imm. al.; Quibula, Beiiguela. Lisbon Museum [P.]. 
skull. {Cotype of species.) 

A-e. 3 (^ ad., 2 Ruwenzori East, Ruwenzori Exploration 
ad. sks.; 5000-.5500^ March, Coium. [F.]. 
1906 (/?. K. Dent). 
Euwenzori p]ast, Riiwenzori Exploration 
o500' ; March, 1906 Couim. [P.]. 
{D. Carruthen). 
Ruwenzori East Ruwenzori Exploration 

(fl. E. Dent). Comni. [P.]. 114. 

Kodja, Gaima Alexander-Gosling Ex- 

Range, N.E.Congo, peditiou [P.]. 
3° 30' N., 29= E. ; 
26 Aug. 1906. 

Measurements of E,ousettus lanosus and angolensis. 



2 2 ad. sks.; 



6 ad., 6 
imm., 2 2 

imm., (5' 
pull. al. 


2 imm. 
sk.; skull. 

if. lanomis. \ R. angolends. 

MiN. Max. 

MiN Max. 

Forearm I 885 

PoUex, c.u. 34 

2nd digit, metacarpal j 37'5 

,, 1st phalanx ■ 8'8 

2nd-.Srd phalanx, c. u 108 

3rd digit, metacarpal 59'2 

,, ] st phalanx 40*2 

„ 2nd phalanx 67 

4th digit, metacarpal 68'2 

1st phalanx 30-2 

,, 2nd phalanx 355 

5th digit, metacarpal 54"5 

1st phalanx 26-2 

2nd phalanx 33-8 

Ear, length from notch 20'2 

,, greatest width, flattened 14 

Eye to tip of muzzle 

Tail 15-5 

Lower leg , 39 

Foot,c. u 23-5 

Skull, total length to front of preraax ! 43'7 

,, width of brain-case at zygomata ! 17'1 

,, zygomatic width | 24'8 

,, postorbital constriction I 93 

,, interorbital constriction i 7'7 

,. width acrofs m'-, externally 1I'2 

„ width across canines, externally 8"7 

,, palation to inci.sive foramina [ 19'2 

front of orbit to tip of nasals i 15'7 

Mandible, length 32 

Upper teeth, c-m'-' 14'3 

Lower teeth, c-in, I 16'2 
































































3. BONEIA, Jcntlnl: 


1879. Boneia, Jentink, Notes LeAfdvn Mm. i. p. 117 (April, 

1879) .' B. biden?. 

Diagnosis. — Closely allied to Housfttas, with wliich it accords in 
most of its cranial and dental and practicallj- all external characters 
(second digit clawed, a short tail, membranes from sides of back), 
but differing in the following particulars : palate much broader 
anteriorl}', premaxilla; separated in front, upper and lower canines 
excessively heavy at base, lower canines directed strongl}- outward, 
inner pair of upper incisors lost (at least in adults), outer pair of 
lower incisors larger than inner, crowns of molariforra teeth flatter. 
General size (of the single species known) as the larger species of 
llousettaa: forearm about 95 mm. 

Fig. 5. — Buncia bidcna, (^ . Meiiado. No. \)~.l.'2.i) (type of 
B. iiiciiadnisk). \, 

>Skull (tig. 5). — DifTcring from that of Jioastttu^ in the two clia- 
racters referred to in the diagnosis above ; in all other respects 

5f? EONEIA. 

perfectly Rousetlinc. In liouscltus tlic breadth of the pnlat.e 
( palatal plates of niaxilhe) anteriorly between the cingnla of the 
canines is lews than, or at most equal to, half the breadth between 
m'^-ra-, the maxillary tooth-rows therefore very conspicuously con- 
verging in postero-anterior direction, and the rostrum, in ujiper 
view, distinctly tapering anteriorly, the breadth across the outer 
surfaces of the canines being much less than the lachrymal breadth. 
In Boneia the palatal breadth between the cingula of the canines 
is about two-thirds the breadth between m"-m", the maxillary 
tooth-rows not conspicuously converging postero-anteriorly, the 
rostrum in u])per view, owing both to the greater distance between 
the canines and to the unusually heavy sockets of these teeth, not 
distinctly tapering anteriorly, the breadth across the outer surfaces 
of the canines subequal to the lachrymal breadth. Premaxilla? 
separated anteriorly by a space of about 1 mm. 

Deflection of brain-case greater than in typical Rovsettus, nearly 
as in Steiionycteris {liousettvs lunosus, p. 49, fig. '•'>), alveolar lino 
if projected backward passing through supraoccipital near upjier 
margin of foramen magnum. Occiput not tubular (comparo 
Ptn-opus). Length of rostrum much greater than lachrymal 
breadth ; front of orbit above middle of m'. Palate, apart from 
greater breadth iu front, quite as in Mmtsetfus, postdental portion 
not long as in Pteropus. Premaxilhe perfectly similar in shape to 
those of Eidolon and Roiiseitus, narrow, u])per extremity curved 
forward and tapering to a point. Tympanic unknown. Sagittal 
crest low, scarcely raised above level of brain-case ; postorbital 
processes as in llousettus. Coronoid low, much sloping, rather 
similar to that of Stenonycteris, though still lower and somewhat 
broader antero-posteriorly ; condyle of mandible below level of 
alveolar line. 

Dentition (fig. S).-.-!--^!'-^^-'"'— x2 = 32. p' and m=, 
i^i,cp,p3p,m, m,m3 

Pj and 51)3 reduced. Dilferential characters, as compared with den- 
tition of liovsetfus, see diagnosis of genus, siiprci. 

i^ small, subterete, crown very slightly differentiated from shaft, 
cutting-edge rounded off; premasilla projecting on inner side of i'^ 
(see fig. 5), suggesting the possibility that a rudimentary i' may be 
present in young individuals, i^ similar in size and shape to i"; 
i„ scarcely differing in general shape, but higher and considerably 
larger, crown area twice or twice and a half that of i^. Upper 
canine long, very broad f antero-posteriorly) at base, compressed 
from side to side, its socket conspicuously projecting on side of 
rostrum ; anteroposterior basal diameter subequal to length of ji' 
(in 7^o«'4W^?/.s about two-thirds the length of p''); profile of front 
margin of crown straight ; anterior surface of crown marked with 
a broad and deep vertical groove : a similar deep but narrow 
vertical groove on hinder surface of crown ; cingulum narrow. 
Lower canine very heavy at base, crown slanted strongly outward. 

p' slightly larger ill crown area lliaii i^ p, about twice tlie bulk of i,. 
p^-in* and p^-m., similar to corresponding teetli of liouscttufi, but; 
crowns Hatter, longitudinal ridges lower; outer and inner ridge of 
]>' and Pj less completely united anteriorly than in liousettjis ; m" 
smaller, scarcely one half the area of m' ; m, little more than half 
the bulk of m^ ; ra^ similar in size to p,. 

Falate-ridijes. — 4 + 3 + ? (formula derived from Jentink's descrip- 
tion, I. s. c). 

Wings. — Second digit clawed. Lateral membranes arising from 
sides of back, separated by a space about 15-17 mm. in width, and 
inserted posteriorly on back of first (or junction between first and 
second) metatarsal. Relative lengths of metacarpals and phalanges 
essentially as in Stenoniidans (Bousetfy.s lauosns, see p. -0). The 
subjoined wing-indices are calculated from measurements of only 
two specimens : — 


,, Pollei 

2nd dif:it. 

3rd digit. 

4th digit. 

5th digit. 

For..arm. ^°"«^- 


1st '2-3 

671 461 




lat 2nd 
ph. ph. 

357 441 


1st ' 2nd 
ph. ph. 

276 ' 3 44 


1000 420 


121 126 

Tail. — As in Roiisetfus. Shorter than hind-foot ; basal portion 
conneclcd with underside of interfemoral by its dorsal integument, 
tip freely projecting. 

liinu/e. — Celebes ; as yet only known from the northern portion 
of the island. 

Affinities. — Boneia is closely related to Enuseitus, but decidedly 
more specialized. It differs chiefly (and almost only) in certain 
characters of the canines and incisors, which again have effected 
some modification of the front of the rostrum and mandible. As 
pointed out in the description of the genus, both the upper and 
lower canines are unusually heavy, and the lower canines slanted 
strongly outward (see front view of rostrum and mandible, rig. o, 
p. 55) ; the latter character necessitates a greater distance (broader 
jyalate) bet^veen the upper canines; the breadth of the rostrum 
anteriorly is further increased by the laterally projecting sockets 
of the upper canines. The widening of the space between the 
upper canines has forced the rami of the prcmaxillne from each 
other anteriorly (in all typical species of lioKsttttis. as in most 
genera of Megachiroptera, the premaxillie are much more firmly 
united with the maxilliC and nasals than anteriorly with each 
other). It appears unlikely that the degeneration, or at least in 
adults complete suppression, of the inner upper incisor has had 
anything to do with the separation of the premaxilhu ; though i' is 
missing, its portion of the premaxillre is preserved. The heavy 


lower canines and somewhat enlarged outer lo\\'er incisors make 
the sympliysis of the mandible distinctly broader and longer (antero- 
posteriorly) than in Bougeitu,^. The tip and trenchant hinder edge 
of the lower canines has carved a deep vertical groove in the front 
face of the crown of the npper canines. The heavy canines and 
anteriorly protruding lower jaw render this portion of the skull and 
dentition somewhat bulldog-like in appearance. The cheek-teeth 
are relativeh" weak (longitudinal ridges unusually low), and, in 
accordance with this, the coronoid process low and much sloping, 
the deflection of the brain-case greater than in any species of 
Eousettus, except the weak-toothed E. lanosus. — The separation of 
the premaxillae might suggest aflinities with Eidolon, the somewhat 
enlarged i., with Pleropus ; but in no other characters does Boneia 
show any leanings whatever to these genera. 

Hisloiv/ of c/emis in literature. — .Secondarj' references: Dobson, 
Hep. Brit. Assoc, for 1880, p. 174 (near Eousettus) ; Winge, E Mus. 
Lundii, ii. pt. 1, p. 59 (1892 : near Eousettus) ; Matschie, Mega- 
chiroptera, p. 69 (1899: near Eousettus); Miller, Fam. & Gen. 
Bats, p. 61 (1907 : near Pteropns), 

1. Boneia bidens, Jtntlal-. 

Boneia bidens, Jcntinli, Xoteg Leyden Mus. i. p. 117 (April, 1879: 
Bone, N. Celebes) : Dobson, Eep. Brit. Afsoc. for 1880, p. 174 
(1880) ; Jentinl; Cat. Ost. Mnmm. p. 264 (1887 : Gorontalo) ; 
id., Cat. Sijst. Mmnm. p. 152 (1888 : Gorontalo ; Bon^) ; Floiver 
cV Lydehker, Mnmm. p. 653 n891) ; Trouesscni, Cat. Mamm. i. 
p. 85 (1897 : Bone) ; Matschie, Mef/achir. p. 69 (1899 : Bone) ; 
Troiiessart, Cat. Mamm., Suppl. p. 60 (1904: Bone): Willink, 
Nat. Tijd. N,;derl. Ind. Ixv. p. 275 (1905 : Celebes) ; Miller, 
Fam. ^'Gen. Bats, p. 62 (1907). 

Boneia nienadensis, 2^homa.i, Ann. S,- Mag. N. H. (6) xviii. p. 242 
(1 Sept. 1896: Menado); Troi/e.-^sart, Cat. Mamm. i. p. 85 
(1897: Menado); Matschie, Majacldr. p. 69 (1899: Menado); 
Trouessart, Cat. Mamm., Suppl. p. 60 (1904: Menado); Wil- 
link, Nat. Tijd. Nederl. Incl. Ixv. p. 275 (1905: Menado); 
Miller, Fam. ^ Gen. Bats, p. 62 (1907). 

Forearm about 95 mm. 

Ears. — Similar in size to those of Eousettus leacJii, but much 
more distinct!}' attenuated above, tip narrowly rounded ofl' ; anti- 
tragal lobe small. 

Fur. — Short ; someivhat adpressed on back and underside of 
body, spreading round neck. Upperside of forearm thinly clothed 
with short adpressed hairs for proximal half; femur and adjoining 
interfcmoral furred above ; tibia naked. Approximate length of 
hairs, back 8, nape cf neck and belly 9-10 mm. Least breadth of 
furred area of back 15-17 mm. 

Colour. — d ad. (skin), type of i>. mcuadensis : Back deep Prout'.4 


hrowu, tinged with rus.sot o!i rump. Entire underside, from throat 
to interfemoral, inchiding tlanks, drab with a slight tinge of 
brownish, rather darker on throat and foreneck than on breast and 
belly. Nape and sides of neck golden bufty. A tuft of rigid 
unctuous hairs on each side of neck, deep ochraceous at base, golden 
buft'y at tip ; colour of tufts, in arranged fur, not conspicuously 
differing from that of surrounding hair. Occiput, crown, sides of 
head, and face darker brown than back. 

Sexual differentiation. — Females of this species are unknown. 
They probably differ in having the neck tufts undeveloped or less 
developed than males. 

Measurements. On p. GO. 

Specimens examined. Three males (two skulls), in the collections 
of the Leyden and British Museums, including the tyjie of tlie 
species and of B. menadensis. So far, these appear to bo the only 
specimens known (December, 1908). 

Range. North Celebes : Menado, Eone, Gorontalo. (" Eotie," 
the type locality of the species, is not, as naturally presumed by 
Thomas and all later writers, the well-known Bone or Boni in 
S. Celebes, but a mountain range and river in N. Celebes, very near 
Gorontalo ; sec Bosenberg, Malay. Arch. p. 257, 1878, and P. & F. 
Sarasin, Beisen in Celebes, i. map iii, 1905.) 

Tijpe in the Leyden Museum. 

Boneia hidens, Jentink ; 1879. — Type locality. Bone, N. Celebes 
( Bosenberg) ; type, an adult male, preserved in alcohol, skull in situ. 
The second Leyden specimen, also a male, is mounted, and ticketed 
Gorontalo {Rosenberg) ; skull extracted, very incomplete. 

Boneia menadensis, Thomas ; 1896. — Type locality, Menado, 
N. Celebes ; type in collection. In this specimen the golden buffy 
colour is restricted to the nape and sides of the neck, whereas in 
the type of B. bidens it extends to the occiput ; in all other 
characters the two specimens are identical. In view of the fact 
(which was not known to Thomas) that B. menadensis is very 
nearly a topotype of B. bidnis, there can bo no doubt that the 
difFerence referred to is only individual. 

a. S ad. sk. ; skull. Menado. N. Celebes, Dr. Chas. Hose [C.]. 
3500' ; Oct. 1895. {Type of B. menadensis.) 



Measurements of Jioneia bidens 




Type of 



Type of 

B. menadciiiis. 













10-5 " 




2ncl-3rd phalanx, c. u 

,'lrd digit, metacarpal 

65 1 62 

63 5 62 

34 34 

41 i 43 

6.3-5 1 61 

26-5 26 

32 1 33-5 ^ 

23-5 1 

J 4-5 ! 

24-5 1 

50-5 50-5 


8 i ... : 

1 44 , 

4th digit., metacarpal 

,, 2ud phalanx 

5th digit, metacarpal 

„ 2nd phalanx 


Skull, total length to gnathion 

„ palation to incisive foramina 

„ front of orbit to tip of nasals 


14-2 ,a 
25-5 * ,1 

,, lachrymal width 

io-8 ; 

■ „ width across canines, externally ... 
,, postorbital constriction 

108 1 




„ width of mesopterygoid fossa 

„ between p''-p'', internallv 







Mandible, length 

„ coronoid lieight 

* Estimate. 


4. PTEROPUS, Brisson. 
Pternjnis (pt.), Dobsoii, Cat. Chir, B. M. p. 15. 

17(}'2. Pceropus, Bn's-sv/i, llr</n. Aiiiut., 2 etl. pp. 13, •'■ ' 

lo3-15o Pt. niger. 

1799. Spectrum, Lacepkle, Tubl. Mnmm. p. 15 \_ncc 
Spectriun, ScopoH, 1777, a genus of Lepi- 
doptera] Pt. niger. 

18()(). Eunvcteris, Grai/, P. if. *S'. p. 04 Pt. melanupogon. 

1870. Spectrum, Grai/', Cat. Monl;. ^-c. pp. 99, 100. Pt. niger. 

lb'70. Pselaphon, Gruij, Cat. Monk. S^-c. p. 110 [nee 
Pselaphus, Herbgt, 1792, a genus of Coleo- 
ptera] Pt. pselaplion. 

H99. Serieonycteris, Matschie, Meyaohir. pp. 0, 30. Pt. subniger. 

1907. Desnialnpex, Miller, Fam. ^- Gen, Bafi^, p. GO 

(29 June, 1907) Pt. leucopterus. 

D'Hujnosh. — Basicranial axis distinctly deflected ; occiput tubular: 
length of rostrum nnich greater than lachrymal breadth ; breadtli 
of palate posteriorly .subequal to breadth between canines. Incisors 
r, — '.,; cheek-teeth ^: p' rudimentary, deciduous, p,, m^, and m'^ 
much reduced ; outer cusp of lower cheek-teeth never deeply bifid ; 
no inner basal ledge in lower cheek-teeth (except in Pt. aiielifums), 
and no well developed antero-intcrnal basal cusp in any cheek- 
tooth. Second digit clawed ; membranes from sides of dorsum and 
back of second toe ; calcar well developed ; no tail. Forearm 
86-220 mm. 

Shull (tigs. 6-S). — (1) DiflTerential characters, as compared with 
skulls of EhJolon (fig. 1, p. 3) and liousettus (figs. 2-4, pp. 17, 49, 
i)'!). — Occiput produced backward and downward, as a short tube. 
Tympanic annular, much narrower than in lioiiscttus. Maxillary 
tooth-rows less divei'ging in antero-postcrior direction ; palate rela- 
tively narrower, particularly in its postdental portion : breadth at 
extreme hinder border subecjual to breadth between canines, much 
less than breadth between p'-p'. Ivostrum more conspicuously 
constricted at diastema c-p\ I'remaxillaD heavier, zvgomatic arches, 
ectopterygoid, posttympanic, and paroccipital processes stronger, 
angular portion of mandible more broadly rounded off. 

(2) The typical Pteropine skull (tig. (i). — The skull of Pt. hypo- 
melanus, as showing no ai)preciable specialization in any respect, 
may serve as a paradigm of a " typical Pteropine " skull, and as a 
basis for a brief summary of the principal aberrations from this 
fundamental type : — Deflection of brain-case moderate, alveolar line 
if projected backward passing through middle or upper half of 
occipital condyles. Nostrum not shortened : front of orbit vertically 
above front (or some point of front half) of m'. Sagittal crest 
perfectly developed throughout its whole length, extending anteri- 
orly very nearly to base of postorbital processes; brain-ease much 
constricted in front, breadth of postorbital much less than of inter- 
orbital constriction. Postorbital processes moderate, reaching 


witliiu sliort^. distance of correspoiidini;' small ijrocx'sses ou zygoma. 
Cororioid process moderately strong, somewhat sloping ; coronoid 
Leight of mandible less than length of lower tooth-row, c-ra^, sub- 
equal to c-m'- ; condjle of mandible considerably above level of 
alveolar line. 

Pig. &.^-Ticntpus JnnKmiclanvf. fomesi, ^ . 
'-■'■ ' ■::' ' ". No. 94.7.] 4.2. 

Dai-vel B.iv, X.E, Borneo. 

Tlifi typical Pteropine skull (without, or with only slight, modifi- 
cations) is found in the Lirge njajority of species of the genus, viz. in 
all species of the Pi. Injpomehinvx (except /'/. S)ih,ii;/fr), mar'iannus. 

rTEKoi'is. 63 

Ciiniirps, ritfi(s, melanotus, niel(inoj)ogon (I'ostriun somewhat shorteucd 
ill I't. Jcei/ensis), vampyrus^ alecto, i'onspicillatiix, and neohibernicus 
groups. Leaving all minor differences out of consideration, the 
prii]ci[)al aberrations from this general type may be classed under 
the foHowing headings [(Jj) and (4)] : — 

(3) Modifications of skull in species with weak dentition (figs. 70, 
8 C). — Species with small or excessively narrow premolars and molars 
sho«- invariably all, or most of, the following modifications of the 
skull : — Deflection of brain-case conspicuously greater than usual : 
alveolar margin if projected backward passing through middle or 
Tipper half, or even through upper margin, of sujiraoccipital ; coronoid 
process of mnndiblc weak and much sloping; condyle of mandible 
situated lower than usual, at level of or only slightly above alveolar 
line ; temporal ridges more or less closely approximated, but not 
fused to form a sagittal crest, or not fused throughout their whole 
length ; brain-case much less constricted in front than usual, making 
postorbital broader than interorbital constriction. Extremes of these 
modifications of the skull are seen in Ft. scapalatus, tuoodfordi, and 
jiersonatas (cheek-teeth excessively narrow), Pt. sidmlger, molosahms, 
insrdaris, plueocephalas, temmiiicJci, epidarins, and mncrotl'i (teeth 
smaller than usual). A similar adaptation of the skull is shown 
by the extremely narrow-toothed liouseitus lanosus (fig. 3, p. 49) 
compared with the normal-toothed species of llousettus (cf. also 

Some species with weak dentition show no appreciable, or at least 
no considerable, reduction of the length of the rostrum ; in these, the 
teeth are iiormally spaced or may even be more spaced than usual 
(Pt. scapidatu:^, tvoodfordi). Generally, however, weak dentition is 
combined with a conspicuous shortening of the rostrum and conse- 
quently rather more crowded arrangement of the teeth. In a few 
species the shortening of the rostrum seems to be due chiefly to 
eidargement of the orbits (Ft. temmincli, personcaus, e2ndarit's, 

(4) Short rostrum combined with heavy coronoid process (figs. 7B, 
813). — In the typical Pteropine skull (as shown in figs. 6, 7 A, 
and 8 A, 7'<. hypomelanus) the rostrum is rather long, the front of 
the orbital cavity approximately vertically above the front of m', 
the coronoid process moderately strong and somewhat sloping, tfie 
coronoid height of the mandible less than the length of the lower 
tooth-row, exclusive of incisors. In a small number of species, 
viz. the typical forms of the Pt. pseJaphua group (Ft. pseJaphon 
(fig. 8 B), 2n^06-i«s, tidiercidatus), all forms of the Pt. scunocnsiat type 
( Pt. nauaiensis, moioens-is, anetinnus, fig. 7B), and the typical iorma 
of the Ft. lomhocens'is group (Pt. Jomhofcusis, 4-o/»7rov'»,s), these 
characters are modified as follows : rostrum considerably shortened, 
front of orbit above back, or posterior third, or even middle of \)\ 
coronoid process uiiusually liigh, broad (aiitero-postcriorly), and 
steeply ascending, coronoid height of mandible more than length of 
lower tooth-row, exclusive of incisors. In all of these sj)ecip.s the 
dcntitiim is heavier than usual, and in nearly all peculiarly modified 


iu one or other direction (either by eiilnrgemeiit of the posterior 
basal ledges of tlie molariform teeth, or development of an inner 
basal ledge in the lower cheek-teeth, or tendcnc}- to splitting of 
inner ridges of certain lower cheek-teeth, or reduction of m^ and 
m^, or excessive development of cingulum of canines and incisors, 
or by several of these modifications combined). Compare skull and 
numdible of I'leralojx'.v. 



-A, typical Plei-oijine skull (Pt. hyjjomdn hilh tomci^i,; see text 

p. Gl. B, skull of a species with short rostrum and heavy dentition 

(Pt. cnieliam(f, 7. 1.1. 2;" 6) ; text p. 63. C, skull as modified in species 

with excessively weak cheek-teeth (Pt. icapulatus,&lX>A.2); text p. (iS. 

All figures \. 

The three principal types of Pteropine skulls described above as 
(2), (;i), and (4) must not be considered sharply separated. There 
is, on the contrary, a perfectly gradual transition from (2) to (3) 
(viz. by species showing various stages of reduction in the sizes of 
the teeth, and therefore also more or less pronounced leanings 
towards the cranial charactei's of weak-toothed species), and several 
transitional stages betAveen (2) and (4) (some of the most instructive 
examples are found in the I't. iiselajiliut group ; certain jieripheral 

VTKHol'US. g5 

species of this group, viz. I>t. insularls aud jJi<>-mY/,hah(.^, C"uroli„e 
Islands, and Ft. laicopten'S, J'hilippines, while preserving most of 
the pselaphon characters in skull and dentition, show a remarkable 
decrease in the size of the teeth, and in these species the short and 
heavy rostrum of a Pi.jyseJnphon is combined with a normal or evea 
rather weak coronoid process: similarly in certain peripheral forms 

Fig. 8— A, tjpicnl Pteropine mandible (Pf. hi^pomefanus iome^i qj 7 U '), . 

see text p. 62. B. vnandible of a with short rostrum,' Leav; 

r Q">"o^'f^ ""'^ steeply ascending coronoid process (Pt. psell 

phon 92.2AI) ; test p. 63. C, n>anclible of a species with exces^i/. y 

weak clieek-teeth {Pt. mipiilatiis, ; text p. 63. 

All figures J. 

of the lomhocemls group, viz. Pt. roJricemis, Kodriguez, and molos- 
»i,nt,, Caroline Islands). Hut any essential modifications of the 
skull, other than those briefly referred to abov(s *lo not occur iu 
tlijs genus. 

Dn^titioa (figs. G, f), 10).- I' ^ ' P' ''' P' °^' °^"' x •^-•'4 * r 

1- , 'i'.iCp, p p m, m,, m, '"* • P- 

rudimentary, deciduous ; p„ m^, and m^ much reduced in size. 

* Individual abnormalities in the tooth formula appear to be exceedinalv 
r«re „, all of P/..o;„„., except Pt.,capuk,tus (a secies cbara'tSSv 

d»-ek-..e.h obht.rated), and va.,p;,n,. (cio.,ely ■A\[J io yiy.utca.,. 0?^ 


(it? rri:7;nns. 

(1) Differential characters, as compared with dentition of Eidolon 
<iig. 1, p. 3) and liousettus (tig. 2, p. 17).- — Dentition on the whole 
considerably heavier. Upiier incisors reliitively much larger, with 
crown distinctly differentiated from shaft, i.^ always conspicuously 
larger than, generally about twice, sometimes three, rarely four, five, 
or six times the bulk of, i,. p' rudimentary, stj'lilbrm. generally 
early deciduous. A pronounced tendency throughout the genus to 
enlargement of the cingulum of the canines and development of 
posterior basal ledges in p-\ p', p.^, p^, and m,, these characters 
being obscured onlj' in certain species with peculiarly weakened or 
degenerated dentition. Other characters ess^entially as in FA'lolon 
and lloiisdhis. 

(2) The typical Pteropinc dentition (figs. 6, 9 A, A', 10 A, A'). — 
Upper incisors nearly equal in bulk and height (i" generally slightly 
smaller and lower), crowded or very narrowly spaced ; crown 
distinctly difl'ercntiated. }>osterior basal ledge (cingulum) generally 
obsolescent in i', narrow but distinct in i'. Diastema i"-c wide, 
subequal to transverse diameter of lower canine at middle, and to 
diastema c- p'. Ujqier canines long, siibccjual to ascending branch 
of prcmaxilla^, slender, gently recurved, in certain species practically 
straight ; cingulum always distinct, as a rule prominently developed 
and forming a conspicuous lira oi' ledge at inner and jiosterior base 
of tooth : front face of crown nuirked by a broad ar'd deep vertical 
groove terminating a short distance above tip of tooth, inner face 
by a sharp median keel from cingulum to tij). p' a minute spicule, 
deciduous, p^ slightly shorter (antero posterior extent) than, but 
nearly as broad as, p^ ; ])' subrectaiigular in basal outline, distinctly 
longer than broad ; m' longer and slightly narrower than p' ; m"" 
much reduced, sulx'qual in cross-section to i\ outline of crown 
circular or elongate. Posterior basal ledge of p^ and p' distinct but 
short, marked off ])ostero-e.\ternnlly by shallow notch from base of 
outer main cusp of teeth. 

Dower incisors crowded or very narrowly spaced, in contact with 
or separated by minute space from canines ; crown distinctly ditfer- 

fiOO skulls pxaniinecl by llie writer, and represpnling all species and subspecies, 
only 10 show deviations from llie normal condition ; no less tJian live of these 
skulls are of Pt. scapulaft'S (total number of sliulls examined, 21). three ot 
Pt. gic/anleus (of a total number of 29 skulls), two of Pt. vawpi/ru!^ (15 skulls), 
none of other species. The aberrations found in I't. f/iff'ni/i>iis anti vamptfrus 
are as follows (on those of Pt. scajmlatug see tliis species, p. 40iJ) : — (1) a well 
developed " i.,' is present on both sides in one skull of Pt. vampi/rus (B. M. 
o5.12.26.y0, locality unknown); the right and left i^ are perfectly alike in 
size, larger than but essentially similar in form to i.,. and situated on inner 
side of tooth-row, at base of canines ; (2) uij and its alveolus missing on one 
side: one Pt. vampijrits t'(7«(^)?/r«s (jg. ad., Java, ; (3) ni^ and their 
alveoli missing on both sides: one Pt. f/ifiantcvs (ad., teeth slightly worn, 
uncertain locality.; (4) an " m^," similar i?i bulk to i., of the same 
skull, present on one side: one T't . <p(iaiitet(.'< (ad., teeth somewhat worn 
Bengal, no. ](16.d); nij is in this skull slightly larger thnn usual; (5) " m^ '' 
present on both sides: one Pt. gigantrus (ad., teeth somewhat worn, Kepal, ; mentioned by Dobson, Cat. Chir. B. M. p. Ifi, footnote) : these two 
"m"'" are very similar in size to m- of a normal skull, whereas "m-" is 
noticeably larcer than usual, about half the bulk of m' (the iw.«sibili1y thereforf 
>t*'ing lliul " m- ' ie supernntnerary, and " m ' " homologous with the Moru;al m'- 1, 

entiatcd, :it least in i, ; usunlly a faint median notch in front edge 
of crown of i.^ ; i^ twice or twice and a half the hulk of, hut only 
slightly higher than, ij. Lower canines generally a little shorter 
and more recurved than upper ones, in front view more con- 
spicuously diverging; cingulum distinct or even strong; usually a 
well-marked vertical groove on outer face of crown. Diastema c-p^ 
wide, subequal to antero-posterior basal diameter of upper canine. 
Pi much reduced, once and a half or twice the bulk of, but scarcely 
higher than, i,, permanent ; crown subcircular or somewhat elongate 
in outline, crushing surface concave ; nearly always situated closer to 
canine than to p^. p^, p^, and ni^ subequal in cross-section, longer 
tlian broad ; m., smaller than m^ ; m^ much reduced, subequal to i,^, 
outline circular or somewhat elongate. Posterior basal ledges of 
Pj and ]\ as in corresponding upper premolars. 

A typical Pteropine dentition, similar to that described above or 
differing only in minor details, is found in about half the number 
of known species, viz. all forms of the Ft. Jn/ponichnnis (except 
Ft. suhniyer), viaritninus. canleejjs, rufiis, melunotas. and melanopor/on 

(3) The principal modifications of the typical Pteropine dentition 
may conveniently be epitomized under the following headings: — 

Upper incisors.— General size larger and cingulum conspicu- 
ously broader than usual, forming a noticeable shelf posteriorly, 
in all species of the Pt. pselaphon group, viz. Ft. pselaphon, pilosus, 
tuberculalus, leucopterus, 'mstdaris, and phaoirphahis. (Compare 

Lower incisors. — In most species i„ is about once and a half, or 
twice, or twice and a half the bulk of ip but in some species the 
disproportion in size is considerably greater. This is sometimes 
due chiefly to a reduction of i,, making this tooth only about \-^ 
of i„ : Ft. Jdiiihoceii^sis, solitarius, rodnccnsis^ and inoloxsit'iis. More 
frequently, however, to actual enlargement of i.^, sometimes com- 
bined with a slight reduction of i, : Ft. iiawaiettsis, mmoensb^, 
anetiatiHS^ psel((phon, p'dosus, tuJiercidati's, and letwopterus ; in some 
of these species i.. is about four or five times tlie bulk of i^, 
(Compare Pteralope^v.) — In Pt. srap^datus and woodfordl all incisors 
are reduced in size. 

Canines. — As a general rule, weak cheek-teeth (no posterior 
basal ledges, or general size of teeth conspicuously reduced) are in 
this genus combined with longer, slenderer, and straighter canines 
with narrow cr even somewhat ill-dctined (but never completely 
obsolete) cingulum, whereas heavy cheek-teeth (strong basal ledges) 
are associated with rather shorter, stouter, and more distinctly re- 
curved canines with strong, sharply detiiicd, shelf-like cingulum. 
In species with very broad cingulum of the canines, the edge of the 
cingulum sometimes shows a pronounced tendency to subdivide 
into a row of small, round, more or less incomidetely separated 
tubercles: Pi. sumoeni^is, andianuK, psehtphon. pilosus, I, uropientjf, 
tubfrculatiis, iiig}dari.i, plKTocephalus (cominro with this Pfernlopex). 
Tn Pt. liiohts.iitiHx the crown of the canines is more strong!)' com- 
pressed fi'nm ^ide to -.idc ihnn ii^u;il. Tlic vertical gro<ive on thtj 


front face of the upper canines is shallow or soraetiraes oltliteiafod 
ill Pi. natidis and livwgsicmei. Pi. hihercuhit^i''; is unique in the 
genus in having a small', but perfectly well-marked, cusp-like pro- 
jection on posterior trenchant edge of upper canine above middle 
of tooth (compare Pterahpe.v). 




Jig. t). — p', p^, ni', profile of left row, pulate view of right row. All figures |. 
A, A', typical Pteropine dentition {Ft. hypcmelamu hiteus, imma- 
ture, B, B', heaTj dentition, strong piisterior basal 
ledges {I't. ])selajilion,&d\i\\,^^1.2A.\). C, C, teeth short (antero- 
posteriorly) and broad, posterior basal ledges practically obliterated 

» {Ft. nenhibernicus, immature,, type of Ft. corenia/Ka). D, D', 

degenerate dentition (Ft. svaptdafus, adult, 

pi. — Eather larger than usual, sometimes equal to or slightly 
laro-er than i , and with crown distinctly differentiated in Pt. pilostui, 
leucopterus, ncnuciiensis, samoensis, anetianus, lomhocensis, and soli- 


p . Vnusually large in Pt. nawaiensis, scvtoeiisis, anetlamis, 

psdophoyi, jvlosug, tulermlatus, and leucopterus. This peculiarity 
is in all of these species associated with an enlargement of i.^. 
(Compare Pteralopex.) 

Posterior basal ledges of molariform teeth. — Species with typical 
Pteropine dentition possess short or moderately strong posterior 
basal ledges in p\ pS p^, and p, ; not infrequently the ledges are 
also more or less distinctly perceptible in m' and m,. From this 
general type the modihcations go in two opposite directions: — 


Either towards complete or iiearlj- complete oliliteration of the 
ledj;;es, as in the s[)ecics of the l^t. </ii/antei(x (five species), alecto 
(four), cimspicillatus (three), neohiheniirux (two), and macrotis groups 
(three), though in young teeth of some of these species a slight 
remnant of the ledge is occasionally detectable. Or towards an 





rig. 10. — p,, ]^^^. in,, m.^. profile and upper view of left r)w. All figures \. 
A, a', typical Pteropine denti'ion ; posterior basal ledges moderate 
{Pt. hiiiKiiiielanux lutcus, iminatiire, B, B', posterior 
basal Ifdges heavy, separated postero-externally by a deep notch 
fmm base of outer main eusp ; note in tlie upper view tiie deep 
transverse groove in the inner (in the figure right) longitudinal 
ridge of p, and ni, in front of middle of ridge {I't. pselaphoii, adult, (-", strong ])osteiior bnsal ledge of p^, nii, and in^ coii- 
tiiiueil forward along ba.>ie of inner (right) side of tooth {Ft. aneti- 
anu.-'. adult,';r)). R, D', teeth short (ant«ro-posteriorly) and 
brotid ; piisterior bisnl ledges practically obliterated (PL reo- 
hiliernii-iis. immnture, 8().7.9.1. type of I'f. rvroiianis). E, E', de- 
geuerat*.' ileutitioii (Pt. scapuUilus, adult, 

unusual strengthening of the ledges, as in Pt. rodrirensis, nawai- 
msis, sarnohms, anttliniu^, innuJdri.i, phceocephaJus, pselaphon, 
pih.tus, and t'lhercnlntus ; in most of these species the ledges are 
■well-marked also in m' and nij. and not infrequently indicated in 

Inner hasal lodge in lower molarit'orm teeth (tig. 10 (V). — I't. 
uiietianus is unique in the genus (but similar to Acerodon) in ha'^iug 

70 rrEKOPCTs. 

a bi'oarl, well-defined inner basal ledge iu p,. nip and ni^. this ledga 
being in reality (quite as in Acerodon) a direct continuation of the 
posterior ledge along the inner side of the teeth. (_)n close exami- 
nation a very faint trace of an inner ledge in the lower cheek-teeth 
is sometimes perceptible in the related Ft. sanwensis and 2^iiosits. 

Traces of antero-internal basal tubercles in p', p', p,, and p^. — In 
a few species the antero-internal cingulnm of p^ and p'' is somewhat 
more differentiated than usual, i. e. more distinctly projecting as a 
narrow ledge : Ft. molossinus and rorh-icensis ; in the latter species 
the rim of this narrow ledge occasionally develops one or two 
minute tubercles. Similarly, the antero-internal cingulum of p^, or 
both p., and p^, shows in certain species a more or less noticeable 
tendency towards differentiation into a projecting ledge : Pf. nuniai- 
ensis, samoensls, (t»ctinni(s, insidaris, and phaoccjihaJ us ; at least in 
J'f. aiieti.anns this ledge develops a minute tubercle. In Ft. psela- 
jifion the antero-internal portion of j)^ is rather sharply marked off, 
by a deep groove, from the inner main cusj) of the tooth. — Compare 

Indications of splitting of ridges of lower molnriform teeth. — In 
the large majority of species the inner and outer longitudinal ridges 
(cusps) of all molariform teeth arc perfectly simjilc. On close 
inspection of p, of Ft. sanioensis and in/osus (allied to Ft. j^selo'phon) 
a faint transverse dejiression in the inner ridge of that tooth is 
detectable, at least in some individuals, suggesting an initial stage 
towards a splitting of tlie ridge into a?) anterior and posterior 
portion. This leads to Ft. i^selophon, in which the inner ridge of 
m^ is always slightly but quite distinctly subdivided into an anterior 
and posterior portion. And finally, in Pt. leiicojitcrns (allied to 
jiseJaphon) not only the inner but, at least in some individuals, 
also the outer ridge of m, and m., are similarly subdivided. — 
Compare Fteralopex. 

m^ and m''.— m.^ is more reduced than usual in the Pt. rayneri 
group (/'('. cor/iKifuf:, rrn/nen, rHhia)nis, Ifu'eUanits, (jrandis, and 
chrijsoprocti(s). Eoth m.^ and m" are reduced in the Ft. loinhocensis 
(Pt. londiocensis, solitavius, rodricensis., and molossinns) and scaj^u- 
latus groups ( Pt. scapidatus and woodfordi) ; in the two latter species 
these teeth are quite rudimentary, and at least in I'r. scupuhtus m^ 
is fiometimes, but nr rarely, lost. 

Modifications of general shape (basal outline), and noticeable 
reduction of size, of cheek-teeth. — The tyjjical Ptrropine shape and 
size of the cheek-teeth lead, on the one side, through practically all 
intermediate stages to the extremely heavy dentition of a Ft. pnela- 
phon, and on die other through numerous transitions to the small 
niid excessivel}- narrow cheek-teeth of a Ft. scapnlatus. Obliteration 
of the posterior basal ledges of the cheek-teeth (a character which 
of itself is perhaps indicative of a beginning degeneration of the 
cheek-teeth) is rather often associated with a distinct reduction of 
the size of the teeth : Ft. r/culdi, epalai-iuf, macrotia, poUocephalux. 
The cheek-teeth are small but not much narrower than usual in 
Ft. niolossiiii's; peculiarly sjiortened, witli rounded corners, in P/. 
papuKHUs and )icohdjcrnici'..^if\gs. 9C', 101) ); and so much shortened 

riKuoPUs. 71 

as lo be ik^ui'Ij' si]u,iris!i in /'/. hucu/iU'nt.s ( [>;iil icuhirly in fJio U])pi'r 
jaw). All the check-teeth are reduced, especially in width, in 
PL suhnvjer. And the extreme of degeneiation is reached h}- 
J't. pgrsonittus, sca/>nh(tHS (figs. 9 1), i)' , lU E, E'), and uwodfon/i, 
in which the cheek teeth are excessively narrow ; a perfect parallel 
to this peculiarity is seen in one species oIl Jiousetiux { li. lanosus) and 
in the Macroglossi ; in the general shu])e of the teeth Pt. personatus, 
aai/mhitus, and woodfordi bear in fact no small resemblance to 
Eonyder'ts sjiehp.a. It should l)e noticed that a reduction of the 
general size of the clieek-teeth (m length only, bieadth only, or both 
in length and breadth) is a character developed independently 
by species of entirely different groups of the genus {Pt. suhnhjer, 
a member of the hifpomelaaus group ; Pt. moJossinus, of the loniboc- 
eu-tis group; Pt. ]ierso)iatHS, of the teinniiucJii group; l^t. .sOTj)M?«<ici> 
and woodfordi, geminate species fbrming a small natural group), 
and therefore far from being indicative of natural relationship. 
The moditication is ])robably in all cases due to adaptation to a 
kind of food which rcijuircs less mastication than that taken by the 
majority of species. 

Fig. 11. — Palale-ritlges, typical Pteropiiie number and arrangeiueMl, funiiiila 
5-|-r)-l-3 (see text, infra) {Pt. hypomelanus tomesi, 94.7. 14.^). 1. 

Falate-riclf/es (fig. 11). — (1) The apparently commonest formula 
is 5 + 5 + 3, i.e. five anterior, undivided, five middle, separated iu 
the median line, and three posterior, approximately wedge-shaped, 
situated near the hindei- border of the i)alate ; tliis formula has been 
observed iu Pt. alecto, inii-tiittws, cJiri/.^ca'chen., tormosi(.s-, 
(jirjaiittus, i/riseus, hi/jwiiieianus, k<'t/eiislx, melaiwpoijon , iiielanotits, 
nttwaiciisig,, iisi laphon, rfii/neri, rudricensis, ruhianus, ton- 
(janus, and v((>upi/ru,t. In some of these species there is a more or 
less incomplete, sometimes nearly fully developed, additional ridge 
between tiie normal ninth and tenth ridges (formula approaching 
5 + 6 + 3: Pi. toni/cnnis. ;/i<ia)iteiis, vainiiip-us, but the same will no 
doubt be found in many of the other species). ISuch species form 
a natural transition to those in which the normal formula seems to 
be 5 + + 3, viz. Pt. anratu.^, rtifus, limhocensis, i^olitarius. In some 
individuals of various species the sixth ridge is only slightly, or not at 
all, separated in tlie middle lino (rormulu nearly or quite (i + 4 + 3j. 

(2) o 4- 4-)- 2 (or 3) pnlafe-ridges have been found in six species : 
A"/, molossinu.-i, ingul.nri!:. phcfocejjhtihis, epidarius,, wood- 
fordi ; in at least two of these (Pt. insidaris anA pJuvocephahis) the 
formula appronches 6 + 3 + 2, owing to the sixth ridge being only 
very alightl}' inteiTupted in the middle. (3) In two species, Pf. pa- 
pwinns and 7uohd)ernicvs, the number of middle divided ridges is 
noticeably larger than usual, formula 5 + 8 + 3. — The palate-ridges 
have been examined by the present writer only in tho tJnrty-one 
species referred to above, but inasmuch as these rei)resent practically 
all groups of the genus, it appears unlikely that any formula essen- 
tially difieront from those given above will be found. 

E:trx. — In most species moderate in length (not reaching eye, 
when laid forward), subtriangular, exposed ; inner margin convex 
from base to tip, outer margin convex in lower two thirds, straight 
or very flatly concave in upper third ; tip obtusely pointed or 
.somewhat narrowly rounded off. The deviations from this general 
type go chiefly in two opposite directions: — (1) Ears considerably 
reduced, half exposed, or so much reduced as to be nearly concealed 
in the fur ; this modiflcation is predominant in two Polj'^uesian 
groups, the Pt. samuensis and ps^elaphon groups, and occurs also in 
a few members of other groups, e. g. Pt. snhnir/er, w<iei\ rodricensis, 
molossinus, and woodfordi; but the line between normal-eared and 
small-eared species is in some cases difficult to draw, the difference 
being one of degree only. (2) Ears long (reaching, or extending 
beyond, eye), outer margin more distinctly concave in upper third, 
ear therefore conspicuously attenuated above, tip acutely or sub- 
acutely pointed ; this modification is characteristic of the Pt. rvfus 
group (except the single Mascarene species), the Pt. vitmpyrufy group, 
the Pt. maootis group, and Ft. caniceps, arf/entatus, and scapuJatys. — 
Pt. livim/stonei is unique in the genus (but compare Pteralopex) 
in having the ears nearly semicircularly rounded oft' above ; it is 
alhcd to species {Pi. melanojtor/on, kn/ensls, ai'uensis) which have 
the ears rather more broadly rounded off than usual. 

Wintjs. — Chief characters : second digit clawed, membranes from 
sides of dorsum and back of second toe. — The line of origin of the 
membranes from tho back varies somewhat in position ; generally 
it is rather nearer the s])ine than the flanks ; in a few species 
(Pt. ivelanopoi/on, papxrinus, ncohihernicus) the membranes arise 
very close together, nearly from the sides of the spine. 

The wing-structure of Pteropus differs from that of the subgenus 
Rousettus (the subgenera Stenonycteris and Lissonjfctens are more 
specialized in this respect ) chiefly in the following four points : — 
(1) pollex relatively longer, index about 440, in Bov,settns about 
340-390 ; (2) second digit with claw (index 740) decidedly longer 
than third metacarpal (index 690), in Ronsettus subequal to third 
metacarpal ; (3) third metacarpal a trifle longer than fourth, fifth 
slightly the longest, while Kov.seitng sliows the more primitive 
cot;dition of having tiie fifth metacarpal tlie shortest ; (4) second 
j)halanx of third digit longer than, rarely subequal to, metacarpal 
of same digit, in llnnnrUnx decidedly shorter than metacarpal. 

Wil.hiu the- genus Plcro/'U.^- the wing-structure is rather uniform, 
the variations chietlj' confined to the greater or lesser relative 
length of the terminal phalanges, and even in this respect the 
variations are rather inconspicuous. .Subjoined the wing-indices 
of Ft. hiJj>omeJanus, calculated from measurements of 56 adult 
specimens representing the eleven known races of the species: — 

2nd digit. 

3rd digit. 4th digit, jj .5th digit. 



0. u. 

1 Mtc. 


120 113 




! , 

2nd| ,,. : l9t 
ph. Mtc. ph 

720 1 670 J408 



40.5 '7I8 

1st ,2nd 
ph. 1 ph. 

302 30S 


437 'i SIO 

Inter femorfd. — In most species scarcely developed, or only a few 
millimetres deep, in centre ; in the species of the Pi. rufus and 
)iiel(niotus groups generally distinct (8-lt) mm.), in the Pi. vampijrm 
group unusually deep (sometimes 25 mm.) in centre. 

Fur. — Typical Pteropino pelage : Pur of back short, distinctly 
adpressed, e.xtcnding on upper.side of lateral membranes somewhat 
beyond their origin from back, breadth of furred area of back 
therefore greater than space between linos of origin of membranes ; 
fur of mantle and underparts generally slightly longer, but difter- 
ence in length often more apparent than real, owing to fur of 
mantle being less adpressed, more spreading than that of back ; 
femur fuired above ; upperside of humerus clothed with short 
adpressed hair; fleshy part of forearm with thinly spread adpressed 
hairs ; tibia naked. Underside of antebrachial membrane, and of 
lateral membrane along outer side of forearm and near body 
between humerus and femur, clothed with short woolh' fur. — There 
is every intergradation from this typical style of pelage (charac- 
teristic of a majority of species), in one direction to species with 
the fur of the back excessively short and restricted to the narrow 
spinal tract between the origin of the lateral membranes {Ft. mda- 
nopoijon, papiuiims, ntohihc micas : aged individuals sometimes 
nearly naked on back), and in the opposite direction through species 
with rather longer fur, extending more or less thickly on the 
upperside of the tibia (most si)ecies of the Pi. rnifncri, Io7nliocensis, 
samoeiisis; pselaphon, and toumincJri groups, and a few aberrant 
species of other groups, viz. Pt. solonionis, hrunneHX, oruatu.i, uuraius, 
niger, and jyoIiocqthah(.^), to species with very long and spreading 
fur {Pt. das)/>nallus, formosus, subniijer) *. 

* It lias been stated in literature that species " provided with long woolly 
liairs extending t>iirkly >ipnn the extremities appear to be coufined to the 
small wind-swept oceanic- islands situuted toward.'* t)ie extreme northern or 
s.mthern limit of the distribution of the family " (Dobson, Cat. Chir. B. M. p. 18 ; 
nee al.M) 'IVmminck, F.^q. Zool. ]i. 6+). In so far a.s tins <t-it.'mc'iit niiffht convey 
the idea that long-furred and haii-y-leej;ed .species arc contiiiMl to the extreme 
iiorllKMn and southern isl.-uuls inhal>iled hy the is n.« well to point out 
that this is I'ar from beiiii: corn-ct. It is true thai on,- of the two species 

Colour. — The most churucteri.slic ieuf.ure is [he clcvelcjuueiil, in 
a majorit}- of species, of a brilliimtly coloured mantle (nape of neck), 
contrasting with the colour of the back *. liut the character is far 
from being constant ; the mantle shows any colour tinge, from light 
yellowish buff or even cream-butf ur whitish, through any shade of 
buff, tawny, russet, or chestnut, to dark brown, seul-brown, or 
blackish. liy darkening of the normal biiglit colour the mantle 
may become similar to the back ; if at the same time the colour of 
the back is lightened, the mantle may even become much darker 
than the back {Pt. aruensi:;, some races uf Fl. Jii/pomehinus, and 
others). — The numerous variations in the total aspect of the colour 
of the pelage may be roughly classed as follows: — (1) PeLige 
approximately dark brownish above and beneath (often sjirinkled 

occuiring in the Liu-kiii Islands, viz. Pf. dasymalltui. has long and spreading 
fur extending upon the upperside of the tibiie, and that the lliree Masciirene 
species are all hairy-legged (one of them, Pt. suhuiyer, a s)>ccies hiding 
in caies, also long-furred), and it is probably on these facts that Dobsou 
based his statement. But tlie second species inhabiting the Liu-kiu Islands. 
Pt. loochoensis, has tiie fur rather short and the tibi.T naked above; one of 
the two species known from Aneiteum (New Hebrides) is long-furred and 
)iah"y-legged (7Y. anetianii>i), the other very short-furred and naked-legijed 
{Pt. gcddici); of the five Australia.n species, one is long-furred with the tibia; 
luiusually thickly clothed {Pt. iio'iocephalu)^), in a second the fin* of the body is 
moderate in length and tl)e tibi;e clotlied above {Pf. /iriiimei/.s). while the three 
otiiers are short-furred and naked-legged. If all the hairv-legged species are 
taken together, they cover the Philippines. Celebes, Moluc(';is, Lesser Sunda 
Islands. Bismarck Archipelago, Solomon Islands, Australia, Polynesia, Micro- 
nesia, Formosa, Liu-kiu Islands, and Masearenes. All the naked-legged and 
short-furred species taken together cover jtraotically the whole area inhabited 
by tlie genus, including nearly all the small wind-swept oceanic islands, with 
the onlj' noteworthy exception of tlie Mascarenes. 

* The dilfiirence in colour between mantle and back, so conspicuous iu many 
species of P/croputi, lias been interpreted as an instance of " jjrotecti ve mimicry " 
(in the sleeping attitude of these bats "the brightly-coloured neck and head are 
presented to the view and resemble the extremity of a ripe cocoanut, with 
which the larger species closely correspond iu size," Dobsou, Cat. Chir. B. M. 
p. 17). This explanation, though perfectly natural, perhaps almost inevitable, 
in the golden age of the theories of " natural selection," " protective mimicry," 
&c., would scarcely meet with universal acceptance now. It would seem, in 
this case, no other theory is i-equired than the very simple one : similar causes, 
similar effects. The real fruits and the pseudo-fruits (Pteropi) hang side 
by side, from the same branches, in precisely similar positions, in jjreciselj' 
similar surroundings, subject lo precisely the same ellects of light and shade, 
and so it has been for countless generations of both ; no wonder, therefore, if 
they become similar in colnur. It has been stated (/. .?. c), obviously to support 
the protective-mimicry-theory, that the contrast in colour between mantle and 
back is " much less developed in the smaller species of the genus" (which, "by 
their small size, are less easily seen "); but the statement is entirely wrong ; a 
strikingly bright-coloured mantle or a mantle practically similar in colour to 
the back are characters equally common in species of all sizes, from the very 
smallest, through the medium-sized, to the very largest. Also it would seem 
that, if Nature had given these bats a protective colour, then she would, so 
to say, have given with one hand and taken back the benelit of her gift with 
the other; the colour of some species may render them not too easily detect- 
able for the untrained eye, but at the same time all species have so strong an 
odour that any of their enemies, be he man or animal, easily detects ihem with 
closed eyes ... , „ ,, „,,,» .„■..,... .-• ■ ,1., ,.^. .^, .-.n, ,„,,. 

with greyish hairs, pariicuhuly on uuclt'i'inirU), and viLh l)ri^htcr- 
<;olo\ireii niaiitle (many spocies. particularly smaller, hairy-legged 
forms, predominant style e.g. in the Pt. samoensis group); (2) back 
blacki^ih or brownish (often thinly sprinkled with greyish hairs), 
mantle bright-coloured, underparts paler than back (many species, 
style purest in Pt. rufus, girinnteus, and their closest allies) ; the 
pale colour of the underparts may be restricted to the centre of the 
breast (Pt. melanotus and others), or wholly replaced by bbickish 
or brownish {Pt. vam/ii/rns and othors); the normally very incon- 
spicuous pale sprinkling of the back may be greatly exaggerated, 
making the colour of tlie back nearly hair-brown (Pf. 7iiadicits) or 
even silvery greyish (some races of Pt. /u/por>ieJ(iina<, Pt. arnensii) ; 
(3) blackish above and beneath, generally more or less thinly 
sprinkled with greyish hairs, and with strongly contrasting yel- 
low ish-bulF mantle (this stylo is found only in the eleven species of 
the Pt. mariannus and conspkillatn^ groups, and in Pt. macrotis and 
epulavius) ; (4) uniform blackish above and beneath, with or with- 
out some trace of a "tippet" (Pt. modigJiani, nafalis, some races 
of Pt. I'ainpi/rus, Pt. aJccfo, gotddi, lii'iny&tonei).- The upperside is 
tricoloured (pale mantle, dark hack, pale rump) in most of the 
species of the Pt. rai/tieri group inhabiting the Solomon Islands 
( Pf. rai/»eri, ruhidjius., Inrelhiims, gra»dis) and in a related ^oluccan 
species (Pt. chri/sojjroctus). The whole of the fur of the body is 
unusually pale, nearly silvery buff or silvery whitish, in Pt. tem- 
tniwlci, persoiuttus, and capistratvs. One species is nearly uniform 
buffy above and beneath (]'t. Iceijeiisis). Pt. persomitus and capi- 
stratus are unique in the genus in having the head marked with 
sharply pronounced dark brown stripes on pale ground (compare 

Sexual differentiation. — Canines in males of nearly all species 
longer and heavier than in females. I'ur of mantle in most species 
which show no other external sexual differentiation more rigid and 
unctuous in males, sotter and more spreading in females (nearly all 
species of the Pt. hypoixehinv^, mariannus, rufus, melanotus, ntelano- 
jtogon.alecto,conxpicillatus, a nd/ifo/aAeTMtoiis groups, but the character 
is not e(]ually conspicuoiis in all of these species) ; in some of these 
the concealed base of the fur of the mantle is dark brownish in 
females, contrasting with the light yellowish-lnifl: tips of the hairs, 
whereas in males the fur of the mantle is light-coloured to the 
extreme base (h. mariannus and conspicillatus groups). Males of 
certain species have a well-developed rigid, generally richer-coloured 
tuft of glandular hairs on each side of the neck, while these tufts 
are either entirely wanting or indistinctly differentiated in females 
(Ft. hruntiei'.s, cognaius, rai/neri, ruhianus, lavellanu-s, grandis, 
lomhocensis, soliiarius. giqanteus, ariel, li/lei, epulanus, macrotis, 
scapulntus, vinodfordi). — l)obson's statement (Cat. Cliir. pp. 17, 49, 
54) that the mantle of the female is generally " darker or less 
brilliantly coloured than that of the male" must have been based 
on insuHicicnt material ; the individual variation in the colour of 
the mantle is in some species considerable, but in no species 

exftmined Ij}' the present writer is there any appreciuble sexual 
ditl'erence in the cvposed colour of the mantle. 

Vi"*/))*/^.— Malagasy region generally, but not extending to the 
continent of Africa : Oriental region ; Austro-Malaya ; North and 
East Australia (not known with certainty from Tasmania) ; western 
Polynesia, north-west to Volcano, Bonin, 8. Liu-kiu Islands (not 
Japanese main islands), and Formosa, east and south-east to 
Caroline, Sta Cruz, Fiji, Tonga, and Samoa Islands. See geo- 
graphical review of species, infrci pp. 79-86. 

Affinities. — Pteropus has originated from a Itousettus-\i\e type. 
It comes nearest the living species of Itouseftus in the general 
shape of the skull, and the dental formula is the same in both 
genera. In at least one point Pteropus appears to be slightly more 
jirimitive than Jiousettus ; the t\mpanic is rather nearer in form 
(narrow, annular) to the generalized insectivorous type than to that 
of IloHsettns (broader, annular ; compare F/idolon : tympanic still 
broader, with short bony auditory meatus). Already in llousettus 
the occipital region shows a slight tendency to lengthening (com- 
pare HovskUus with Oynoptenis and allied genera) ; in Pteropus 
it is more distinctly lengthened into a short " tube"; the rostrum 
is relatively somewhat longer, the palate narrower, especially in its 
postdental portion, the skull on the whole (in all typical forms) 
more heavily built, with stronger crests, and broader and more 
steeply ascending coronoid process of the mandible. The dentition 
of Pteropus is conspicuously heavier, i.^ enlarged (a slight enlarge- 
ment of ij, as compared with i,, is already seen in certain species 
of Nonsett^is ; compare also lioneia), p' still more reduced and 
generally deciduous (comj)are liovsettus hrachyotis), the cingulum of 
the canines and incisors stronger,' and the posterior portion of the 
molariform teeth generally differentiated into a more or less con- 
spicuous basal "ledge" (in certain specialized forms of the genus 
this ledge is again obliterated, but even then a trace of the ledge 
is often detectalile in the young unworn teeth). Also externally, 
Pteropus is decidedly on a higher level than : the tail 
has disappeared, its wing-structure is slightly less primitive (see 
p. 72), the mnntle generally conspicuously different in colour from 
the back, and the size of the animals generally much larger. Both 
genera are distributed over continental S. Asia, Indo-Malaya, 
and Austro-Malaya ; but whereas Roiisettus extends to Palestine, 
Cyprus, and the Ethiopinn region, exclusive of Madagascar and its 
adjacencies, Pteropus finds its western limit in the Indian Penin- 
sula, including Ceylon and the Maldives, is rather richly represented 
in the whole of the Malagasy region, but perfectly unknown in the 
African continent ; and eastward, liousettus does not extend beyond 
the Solomon Islands, and is unrepresented in Australia, whereas 
Ptero^ms has spread over the whole western portion of Polynesia, 
northward over Micronesia, and southward to Australia. 

Suhdivisions. — The 82 species (1*9 distinguisliable forms) are 
arranged in seventeen groups according to their probable natural 
affinities, viz. :— the Pt. hi/pojiielanvs {Hi species, 26 forms j, )nnri- 

annvx (8 species), caniccjis (3), rafus (5 species, 6 iorms),mehinotiis 
(5 species), melanopogon (4), ratjneri (6), lomOocemis (4), samoensis 
(3), padaphon (G), teumuncld (3), vampyras (5 species, 11 forms), 
«Zerfo (4 species), mnspicillatus ('i), ncohihernicm (2), 9?ia(To;i»- (3), 
and scapulatus groups (2). — From a broader point of view these 
seventeen groui)s may be classed in three primary sections, as 
follows : — (1) The first six groups, the I't. hypoimlanns, marhmmis, 
caniceps, rufus, inelaiiotm, and inelano/togon groups (altogether 
41 species, 52 forms, thus almost exactly half the number of 
known forms), are chiefly characterized by the typical Pteropine 
slcull and dentition (posterior basal ledges of premolars distinct) ; 
the large mujority of species are short-furred and naked-legged ; 
and they cover practically the whole Pteropine area with exception 
of the Himalayas, India, and Ceylon. (2) The second section, 
the Ft. rnyncri, lombocensis, sdmoeiisis, pselapJion, and tennnlncJri 
groups (22 species), accords with the first in having distinct, or 
even unusually strong, posterior basal lodges in the jn-enjolars, but 
the rostrum is always shortened and the dentition modified, either 
by reduction of m^, nr, and i,, or by enlargement of i^ and p,, or 
the whole of the dentition is noticeably (sometimes even excessively) 
weakened; nearly all species are hairy-legged; and they range 
over the whole of Austro-Malaya, Polynesia, and Micronesia, being 
represented also in the Mascarenes ; but are unknown from the 
whole of the Oriental region, except the Philippines (one species). 
(3) The third section, the Ft. vampyrus, alccio, consptcillaius, neo- 
Jiibernicus, macrotis, and icapulalns groups (19 species, 25 forms), 
is characterized by the practically complete obliteration of the 
posterior basal ledges of the cheek-teeth ; the skuU is tyiiical 
Pteropine (except in a few species with degenersite dentition) ; all 
species but one are naked-legged and short-furred ; they range over 
the whole of the Oriental region (being the only section represented 
in the Himalayas, Ijidia, and Ceylon) and Austro-Malaya, as far 
east as the Solomon Islands, extending also to Australia ; but is 
unrepresented in Polynesia, Micronesia, and the Malagasy region. 

Fteropus^ Brisson ; 1762. — Three species were "included by 
Erisson in his genus Fteropus ; the first headed " Fteropm,'' the 
second '■ Pteropus culh riihro;' and the third " Fleropus auricuUs 
patulis;' The first species, as being evidently considered by 
Brisson the Pteropus, must technically be taken as the type of the 
genus, on the tautology princii)le. It is marked in Brisson's book 
with two asterisks (indicating it as a species examined by the 
author himself in the Koaumur Museum), diagnosed " Pteropus 
rufuK aut niger, aiiriculis brevibus aeutiusculis " ( tlie last throe woids 
absolutely excluding Ft. vcmpyrun), and stated to occur in the 
island of Bourbon ( Reunion) ; these facts are evidence that Brisson 
had before him a Ft. niger, Kerr (Ft. vulyco-is, E. Geoff. J, not a 
Jt. vampyrus. But, as might be expected, Brisson places in the 
synonymy of the lieunion spc(;ies also references to Seba's Can-is 
I'olaits Ter)iutatin.i orientalis (which is Ft. vampyms) and Linne's 
VespertUio cnuda nvlht (l)ased on Seba), and therelore gives as 

liabitat not only Ueunion lait al&o Ternate. — Owiuo to a uiislakcn 
identification of Brisson's first species of Fteropiis, tlie type of this 
genus has hitherto been given as Fi. vnmpyrus (Merriam, Science, 
(n. s.) i. p. 376, 1895 ; Miller, Fam. & Gen. Bats, p. b'o, 1907: 
probably by a slip Matschie gives on p. 6 of his ' Megachiroptera ' 
the type as Pt. conspiciUatus, on p. 12 && Ft. cdano = Ft. vampyrus). 

The genus Fteropus as understood by Dobson in his ' Catalogue of 
Chiroptera ' (187S) corresponds to the genera Ftcropvs, Acerodon, 
and Styloctenium of the present catalogue. The genus as hero 
defined is equivalent to Fteropus and Desmcdopex in Miller's 
' Paniilies and Genera of Bats ' (pp. 56 and 60, 1907). 

Spectrum, Lacepede ; 1799. — Type (only species), '^ Spectrvm 
vampirus." Linne's Vespertilio vamj^yims (1758) is a composite 
species, including Ft. vampyrus, mger, and rufus. Of these, 
Ft. niger uas apparently the only species known to the early 
French mamnjologists from autopsy (see Brisson, /. s. c, and Bnffon, 
Hist. Nat. X. p. 55, who distinguishes only two species of Fruit- 
bats, the Koiissette, which is i;nquestionably Ft. niyer, and the 
Rongette, which is Ft. suhmyer, l)oth from the Mascarenes), and 
maj' therefore be fixed as the type of Lacepede's Spectrum. — In a 
revised reprint of Lacepede's 'Tableaux methodiques,' published 
in 1802 in the Didot edition of Buff'on (Quadr. xiv. p. 18S), Daudin 
includes two species in the genus Spectrum, S. vampirus and 
S. ruhidum ; the former is Ft. niger, the latter Ft. subniger. 

Euiiycteris, Gray; 1866. — Type (only species), '-Fteropus 
phaioyjs," Tenim. Temminck's Ft. pJia-iojis, 1825 (Mon. Mamm. i. 
p. 178), is Pt. rufus, E. GeolT., J 803 {Ft. edwardsi, E. Geoff., 
1810); Temminck's Ft. pltaiopis, 1837 (Mon. Mamm. ii. p. 65), is 
Ft. melanopogon, Peters. From Gray's diagnosis of Euvycterts 
(only important characters: uings attached to the vertebral line, 
the part over the back bald, or nearly so) it is clear that the 
" P/. pliaiops'' which he proposed to separate generically from 
Ft('rop)us is Ft. melanopogon.- — Eunycteris has been considered a 
synonym of Fteropus by all authors except Jfatschie (ilegachir. 
pp. 6, 11 ; 1899), who revived the name for a subgenus of Fieropnis 
including four (in reality three) forms, Vt. melanopogoti, papuanus, 
<hgfner [^neoJnberaicus I, and neohilernicus. I tail to see any 
chaiactcrs that bind these three species together in contiadis- 
iinction fo all other species of the genus, except the narrowness 
of th« furred area of the back. Ft. jiajiunnus and neohilernieiis 
are essentially -ditl'erent fiom Ft. 'inel(inopot,on in dentition, and the 
latter species has no closer known allies than Ft. aruensis and 
keyensie, which by Matschie were referred to the subgenus Fl(ropus. 

Spectrum, Gray ; 1870. — Founded by (jiray independently of 
Lacepede's Spectrum, with which he was ajjparently unacquainted. 
Separated from Fteropus on account of its round head, narrow face, 
and small hidden ears. Five species : S. vulgar^ [Ft. niger; type 
by subsequent designation: see Matschie, Megachir. p. 6 (1899)], 
rnbricolJe [Pt. suhnir/er^, dasymallum, aneiionum, and levcnpierum. — 
Name revived by Matschie (/. s. c.) for a subgenus of J'teropus 

iiK-luding twenty tlve species. It is a purely aititicial constellation 
of species, indefinable as a genus or subgenus. 

Fselaphon, Gray; 1870. — llests solely on the combination 
" P.-itlaphon ur&hnis" occurring in Gray's Catalogue [l. s. c). Eut 
that this is a slip I'or Fteropus ursinus is beyond all doubt : 
Fsekijihon ursimiit is no. 23 of the genus Pteroptis, preceded 
and followed by many other species all of which are styled 
" Pieropus " ; a genus Psdaphon does not occur in Gray's synopsis 
of genera of the family Pteropodid.e in the same book (p. 99); and 
in the systematic index to the book (p. viii) the species is registered 
as Pteropus ursinus. 

Sericoni/cteris, Matschie ; 1899. — Subgenus of Pteropvs ; type by 
• ■riginal designation, Pt. ruhricoUis [Pt. subnif/er], Si)eeies sis : 
J't. nibricollis, teniminc/ii, personatus, capistratus, molossinns, tcood- 
fordi. It is difficult to understand whj' these species were placed 
together in a distinct subgenus ; the brief diagnosis given by the 
author does not touch any character of importance and does not 
hold good on any point ; the six species, representing, as the}' do, 
four different groups of the genus, have in fact little more in 
common than their unusually small size. 

iJesmalope.r, Miller ; 1907. — The type and only species, Pt. leuco- 
pt/THs, differs in no character of generic importance from Pt.p>seliiplion 
(see Ann. & Mag. X. H. (8) Vi\. pp. 1^13-218, 1 Feb. 1909). 

Geor/ntphical Distrilmtion of the S^iecies and Subispecies 
of Pteropus. 

I. Midaijamj Reyiou. — 8 species (9 foims), re{)rescnting four 
different groups of the genus, \iz.: — (1) Pt. nifus, comorengis, 
scycliellensis, nldabroisis, niijer (all of the Pt. rufus group) ; (2) 
J't. Jivingslonri (Pt. itiehinopO(/on group) : (3) Pt. s^ibnigcr {Pi. 
Iiypomehtiius group); (4) Pt. rndncensis (I't. loinbocensis group). 

All the species arc allied to Indo-Malayan or Austro-Malayan 
forms, none to species inhabiting India. The absolutely pre- 
dominant type of Pteropus in the ifalagasy region is the Pt. nifus 
group : it is the only group peculiar to the region, and the only 
one distributed over i)ractically all its islands. It falls, however, 
into two sharply separated sections, a Malagasy type, represented 
by four closely related species, Pt. rufus (Madagascar"), Pt. como- 
reusis ((.onioros), I't. sei/chelleitsis (Seychelles), and Pt. aJdabrensIs 
(.ildabra), and a ])eculiarly modified ifascarcne type, represented 
by a single species, Pt. x/r/f/- (ifauritiiis, lieunion). The Pt. rujns 
group has no do.-cr living relatives than the si>ecies of the Pt. 
uichinotus group, all of which are eonhned to t]u'. Andauian-lS'icobar 
chain and its south-eastern contiiniation (Nias, Engano, t'hristinas 
island). — J't. Iivhu/st07in (apparently restricted to Johanna Island, 
(Comoros) is the only Malagasy species of a group represented 
cLsewhere only in the Moluccas, Key, and Aru Islands. Likewise, /'/. 
s«///i(V/f/- ( Mauritius, Reunion) is the only Malagasy representative 
of till' chiefly Austro- and 1 iido-Malayan I't. ItitjionK'hiiimi group. 

80 I'TEKorrs. 

The single species inliabitiug Rodriguez (PL rod riccn sis) is rather 
closely related to I't. lomhoctnsis and noUtarius (Lesser Binida 
Islands").- It is a remarkable fact thafc in the Mascarenes (the 
small islands of Mauritius, Keunion, and llodriguez) have Leeu 
preserved representatives of three different groups of the genus 
(rnfiis, hypmnelanns, and Jomhocen^is groups), two of which (the 
second and third groups) are iound nowhere else in the Malagasy 
region. Of the three s])ecies, two {Ft. niger, sribmr/er) are common 
to Mauritius and Reunion, while the third is only known with 
certainty from Rodriguez. 

II. Ceylonese, Indian, and Indo-CJiii'ese Svhretjions (including 
Mergui Archipelago, Formosa, and S. Liu-kiu Islands). — 8 species 
(10 forms), of three groups, viz.: — (1) Ft.ariel, fji(juiitnis (f/iyaniena 
and leucocaphahts) , internifditis, hjlti (Ft. vampyrtis group) ; (2) 
Ft. hypouteJanus {coudoreitsis and yeminor)(m),j'oi'mos>is, dasymallus 
{Ft. hypomelanvs group) ; (3) Ft. loovhoensis (Ft. mariannns group). 

The genus is remarkahly poorly represented in continental South 
Asia, in fact chiefly, if not entirely, by western offshoots of Indo- 
Malayan types. Ft. inttrmedcKs (Teuasserim), (liyauteKS (Hima- 
layas, India, Ceylon), and arifl (Maldive Archipelago) are scarcely 
more than slightly modiiied immigrants of the Lido-Malayan 
Ft. vamjiyriis: it is not unlikely that, with much completer mate- 
rial, they will he considered subspecies of l^t. vampyrus. The 
.small Ft. lyJei (Siam, 8aijjon) is rather more distinct, perhaps a 
truly indigenous continental type of this group, as opposed to 
Ft. varnpyrus and its continental ramifications. Ft. Jiy/ionu-laniis 
condorensis (Cambodja, Siam) and yeminoruiu (Mergui Aichipelago) 
are western races of an Indo- and Austro-Malayan species. The 
single species known from Formosa (Ft. formosus) and one of 
the Liu-kiu species (Ft. dasyniaUvs) are represeutatives of the 
Itypouidanus group, m skull and dentition very closely agreeing 
Mith the typical Indo- and Austro-Malayan species of that group : 
the second Liu-kiu species (Ft. loochoensis) is decidedly of Polynesian 
origin, being an only slightly modiiied, extreme north-western off- 
shoot of the otherwise exclusively Polynesian and Micionesian I'i. 
mariannus group ; its nearest xelative is iound iu the Mariannes. 

III. Indo - 2Jid"ya (excl. Mergui Archipelago). — 13 species 
('2'2 forms), of five groups, viz. : — (1) Ft. tytUri, tnehinotHS, niudicus, 
'inodiyVmnii, natalis (Ft. melanotus group); (2) Ft. vainpyriis; ('•'>) 
Ft. hypotntlanus^spenus'iis., ynimus, satyrus, faunulus (Ft. hypo- 
rnchmus group) ; (4) Ft. atenimiis (Ft. alecto group) ; (5) Ft. 
hucojjterus (Ft. pselopihon group). The subregion is easily divided 
into two provinces : — ■ 

(«) Andaman-^'icobar chain and its south-eastern continuation 
(Nias, Engano, Christmas Island). — 9 species, of three groups, 
\iz.: — (1) Ft. fytleri, melanoiiis, ^liadicas, rnodiyhanii, nataiis; 
(2) '•'■ Ft.vampynis"; (3) J't. hyjiomehmus (snhf>\). enr/ariii.i),s(ityrvii, 
faumdiis. — Zoogeograjihically the most inti-i'csting jiarf of the sub- 

rrmiopus. 81 

region. All the fonus einiuu'rated above ^•■/'/. I'diiijn/rn.'! " pcrlKips 
excepted) are peculiar to this proviucu'. The largelj' predoiiiiiiant 
type is the Pt. mflanoUis groui» ; it is' entirely confined to this 
narrow chain of islands, but ratiier closely related to the Malagasy 
Pt. rufus group. The Indo- and Austro-Malayan Pi. hi//>omd<imts 
group has developed peculiar forms in the Andainans (Pt. natjirus), 
Nicobars {Pt. fauiudus), and Eiigano {Pt, hijpomdanus eiKjanus). 
The representatives of the Pi. vmnpi/rus type occurring iu the 
Andamans and Xicobars are as yet imperfectly known. 

(6) Malay Peninsula, Sumatra, Java, liali, Borneo (with 
Natuna and Tambelan Is,), Philippines. — G species, of four groups, 
viz.: — {\) Pt. vampiifiis; (2) Pt. hypomdanus, .yjeciosns, 'niinins ; 
(3) Pt. aterrimns ; (4) Pt. leucopterus. — Although vastly larger than 
the Andaman-Nicobar chain, this province is considerably poorer 
in species. The most characteristic type is decidedly P/. vamjvfrus. 
The range of this species almost exactly covers the ludo-Malayan 
subregioti, only in the extreme south-east it extends beyond the 
limits of this zoogeographical area, to Lombok, 8avu, and Timor ; 
but it has very close relatives in continental S. Asia. Within the 
borders of the present province Pt. vampyms is ditferentiatcd into 
five races, one confined to the Malay Peninsula and Sumatra 
{mahiccmsis), a second to Java {vnmpyrus)., a third to Bali (plutoti, 
extending also to Lombok), a fourth to Borneo and the Natuna 
Islands {nniimc), and a fifth to the Philippines {htiwiisis). Por 
reasons given eise\vhcre it appears probable that Pt.Jiypoindanus is 
of Austro-Malayan origin ; in the present province it has developed 
five very clojely interrelated races ; of the two other species of the 
hypomehtnus type, one {Pt. speeiosus) appears to be confined to the 
province, while the other {Pt. mimus)is common to the Philippines 
and Celebes. Pt. ati'rrimus (Bawean Island, Java Sea) is the only 
representative of the Austro-Malayan and Australian Pf. ahcto 
group. A connection between the Philippine and Microiiesian 
Pleropns faunas is established by the occurrence in the former 
group of islands of a re])resentative of the Pt. pseJaphon group 
(otherwise exclusive!}' Mieronesinn), viz. Pt. Jeucoptcrns (compare 
the spreading of the hypomehinus group to Formosa and the Liu- 
kiu Islands). 

IV. Amtro-Mulnya. — 35 species (38 forms), representing twelve 
groups of the genus, viz. : — (1) Pt. hypamelaiuis, (tdmir(tlitataiii, 
colomis, solo))ioais, mirntiK, (jriseus, jxillidus ; {2) Pi. floLs-otii, 
canicrps, aryentntus ; (3) Pt. melduo/Kiyou, Jceyensis, aruensi!^; (4) 
Pt. coynatiis, r<iyneri, rulnanns, hiveUanvs, yrfnttlif!, cJtnf.iojtnjctus ; 
(.j) Pt. lomhoeensis, soUtai-iHs ; (6) Pt. temminckl, ca pi strut us. perso- 
iinlus : (7) Pt.vampynts; {H) /'Ldlncto, niorio ; (d) Pt. fhryfidiu-hfn, 
ro)i.<tpiclllatus, ociddris ; {\0) Pt. papwanin, neoluhemicus \ (11) Pt. 
mitcroth. epxldrim; (12) Pt. woodfonli ( I't. saijitilKtu.t ^roii]}). 

No faunistic area can comjjare with Austro-Malaya in the 
wealth and multifariousness of its Ptempus population. The con- 
trast between the Indo-Malayan and .\ustro-Miilayan faunas is as 



■well-marked, and the change from the former to the latter as abrupt, 
in this genus of hats as in any group of non-tl\ ing mammalia ; it is 
in this case an almost sudden change from a remarkably poor to 
an extremely rich and varied fauna. The number of species thus 
far known from Austro-Malaya is, as said above, 35, as against 13 
from Indo-Malaja; but it the chain of islands extending from the 
Andamans through the Nicobars, Nias, and Erigano, to Christmas 
Island, with its rather peculiar Fteropiis fauna, were cut off' from 
Iiido-ilalaya, the contrast would become greater still : 35 Austro- 
Malayan against only 6 ludo-Malayan species, or twelve mostly 
richly differentiated Austro-ilalayan against four poorly represented 
Indo'-Mah.yan groups of the genus.— t)f the 'A5 species, 31 are 
peculiar to Austro-Malaya. The four species which extend beyond 
the limits of the subregion are Ft. hiipomelaniis, rnhuus, vampijrvs, 
and consi>inllatiis. Ft. vamp>/rus has very little to do with tiie 
genuine Austro-Malayan fauna; it is an Indo-Malayan type ex- 
tending only to the south-west corner of Austro-Malaya (Lesser 
Sunda Islands). Ft. rninnis is perhaps a Cclebeiin species which 
has spread to the Philippines, rather than the reverse, and Ft. 
conspiciUatus is undoubtedly a New Guinean species Avhich has 
made its way to N. Australia. Only Ft. hiipomelanus is widely 
distributed both in Austro- and Indo-ilalaya, but it has probably 
originated in the former area. 

Of the twelve Austro-Malayan groups of the genus, four are 
peculiar to this fauni.stic area, viz., the Ft. cunkeps (three species), 
rayneri (six), temmincl-i (three), and neoJubernicus groups (two), 
Tliree, of the remaining eight, groups are common to Austro-Malaya 
and Australia, but unrepresented in Indo-Malaya, namely the Ft. 
coiispieillatus (two Austro- Malayan species, one common to New 
Guinea and N. Australia), macroiis (two Austro-Malayan, one 
Australian species), and scajmlatns grou] s (one Austro-Malayan, 
one Australian species). There can be little doubt that at least 
two of these groups, the cnnspkillntus and macrotis groups, are of 
Austro-Malayan, or more strictly Papuan, origin and have spread 
comparatively recently to Australia; as to the third group, the 
Ft. scapulatits group, 'which numbers only two species, one in the 
Solomon Islands {Ft. woonfonli) and one in Australia (Ft. scapu- 
lattis), both with precisely the same peculiar modifications of the 
dentition and skull, its place of origin is doubtful, but the group is 
closely connected with the Austro-Malayan Ft. macrotis type.— Of 
the remaining five groups, one, the Ft. aUcto group, is nearly 
confined to Austro-Malaya (two species) and Australia (one), but 
has spread to the south-east corner of Indo-Malaya (Bawean Island, 
one species); it is probably of Austro-Malayan origin. A second, 
the Ft. melanopof/on group, is common to Austro-Malaya (three 
species) and the Malagasy region (one), a third, the Ft. lomhocensis 
group, to the same two areas (respectively two and ono species) 
and Micronesia (one). Finally, the last two groups, the Ft. Jiypo- 
melanas and vampijntu types, are common to Austro- and Indo- 
Malaya ; of these," the former has probably spread east to west, 

pri;u(»i'us. S;^ 

■R'liereas I't. tuDnpi/rtis is an iiiclo-Mulayau type wliicli has spread 
only to the south-west corner of the ])re.sent area. 

Also the negative features of the Austro-Malayan Pteropys fauna 
must be referred to here. The five jjroups of the genus which 
have no representatives in Austro-Malaya are the Pt. mnriannus, 
samo'ensis, ^'seJaphon, vieJanotus, and i-ufus groups. The two 
former are entirely Polynesian, but one of them, the mariannus 
group, is phylogenetically very intimately connected with the 
Austro-Malayan /t//y;o*»eZa/u<« group, and the other is closely related 
to the Anstro-Malayan Pt. lomhocensk group. The pseiaphon 
group is Micronesian and Philippine, but as lieing closely connected 
with the samoetuiis grou]) leads again back to Pt. lomhocensis. The 
melcmotus and rafua groups are geminate types, the one confined 
to the Audaman-Nicobar chain, the other to the Malagasj' region, 
and related to the hjipomelanus group. 

The two facts, that the Pteropus fauna of Austro-Malaya is incom- 
])arably richer than that of any other zoogeographical area of similar 
size, and that all groups of the genus not represented in Austro- 
Malaya are very closelj' related to Austro-Malayan types, /. e. 
pointing back in their probable origin to Austro-Malaya, lend 
support to the hypothesis that the centre of dispersal of the genus 
was this subregion, or rather an area much larger than, but in- 
cluding, Austro-Malaya and probably excluding Australia. But 
any hypothesis is unsafe, so long as we know very little about the 
physical changes that have taken place in the area between 
the Malagasy region and the Indian Archipelago, and nothing 
about the past history of the genus. 

V. Australia. — 5 species, of five groups, viz. : — I't. coiiKpiciUaius 
(Pi. con.tpicillutu.t group), Pt. poUocephalun (Pt. mairuiis group), 
Pt. goiiJdi (Pt. a/ecto group), Pt. brnnneus {Pt. Iii/poDirhiniis group), 
Pt. srapulains {Pt. xra/ndntits group). 

The small number of species, representing as many groups, is 
evidence that the genus forms no part of the original fauna of the 
continent. The first species, Pt. consjudllatus, is common to S.E. 
New Guinea and Australia and has very close relatives in W. Xew 
Guinea and the Moluccas ; the second, Pt. poVioci'jthalnx, is a j)eculiar 
modification of a New Guinean type. The third species, Pf.^jouhii, 
points toward the Lesser Sunda Islands and Celebes, being very 
closely allied to J't. alccto (Celebes, Sahn'er, Lombok). The fourth 
and fifth species, Pt. hrunnens and srapulotus, have their closest 
relatives in the Solomon Islands. Of the five Australian species, 
I't. •^coptilatns and poliocepJuihis are decidedly the mo*;t peculiarly 
modified, hence perhaps the earliest immigrants into the continent. 

VI. Polynesia. — 18 species, representing live groups, viz. : — (1) 
ft. pdewensis, ijapenxis, tiaJaims, marmnnnx, vdiiih-orensifi, f/eddii-i, 
tiiiii/anus {Pt. ituirlaiinus group) ; (2) /''/. lutwmeiisis. namoi'iisi't, 
unetJanvs {Pt. R(tmo'n\s'is grou])) : (3) Pt. InsuJans, filiaocephalus , 
jisfliiphon, pilosKs. ttiJi.txi'latiis {Pt. pselaphon group); (4) Pt. 

G 2 


oniatus, auratus {Ft. liypomelanus groiip) ; (5) Pt. moJossinns (Pf. 
Joiiihocensls group). 

The relatively large number of species is chiefly clue to the spread- 
ing of a few fundamental ty{)es over numerous widely scattered 
groups of islands, and their consequent differentiation into many, 
but often only slightly differing, forms. The fauna falls naturally 
into three classes of forms — those which are the extreme eastern 
offshoots of essentiiilly Austro-Malayan groups, those belonging to a 
group which, though essentially Polynesian, is also represented in 
the indo-Malayan Archipelago, and the peculiar Polynesian types. 
To the first category belong P<. ornatus (Xew Caledonia) and cmratus 
(Loyalty Islands), only slightly modified representatives of the 
Avidely distributed but chiefl}' Austro-Malayan hypomelonns group, 
which here finds its extreme south-eastern limit ; and Pt. molos- 
st/i?«s (Caroline Islands), a member of a group which now numbers 
only four species, but formerly must have had as wide a distribution 
as the Jiifpomelanns group, since its surviving forms are spread 
over the Carolines, Lesser Sunda Islands, and Mascarenes. To the 
second class of forms l>elong the five speciea of the Pt. pselaphon 
group, inhabiting the Bonin and Volcano Islands { pstlaphon), 
Pelew Islands (piJosus), Carolines (insnhfris and pTuwctphalus), and 
(?) Vanikoi-o (ijfi'x'j-f '(?«<!««) ; this group is represented also in the 
Philippines (I e%i copier us). All the remaining ten species, the genuine 
Polynesian fauna, are clearlj' modifications of only two types, the 
mariannus and samoensis tj'pes. The Pi. mariannus group, seven 
closely interrelated species (with an eighth species, Pt. loorlwensis, 
in the S. Liu-kiu Islands), is distributed over practically the whole 
subregion, so far as inhabited by the genus, from the Pelew and 
Marianne islands south-east to Tonga, Tiji, and Samoa ; it is 
extremely closely connected with the Itypomdanus group, scarcelj' 
more than its Polynesian representative, Pt. admiralitatum forming 
a transition between the two groups. The Pi. samoensis group, 
three sharply differentiated species, is entirely South Polynesian 
(New Hebrides, Fiji, Samoa) ; but the group is undoubtedly a modi- 
fication of the lonihocensis type. 

The subjoined geographical review of the species and sub- 
pecies is based almost entirely on material examined during the 
preparation of this C!atalogue. 

1. Malagas)! liecjion, 

Madagascar. — Pf. rufns rn/iis (northern and central), nifus princcps 

Comoro l.slands. — Pf. comom/sis (generally distributed), livuit/stonei 

(Johanna I.). 
Aldabra. — Pt. aldalircmU. 
Seychelles. — Pt. sci/cheUeusis. 
Mauritius, Reunion. — Pt. niger, sahnic/cr. 
Eodriojiie?:. — Pt. rocJriccnsi^. ... 


2. CnjJonese, Ttitliun, and Lido-C/nnese Suhrcjiom. 
Maldives. — I'f. ariel. 

Ceylon, ludian Peninsula. — Vt . qicjanteu^ giqnntem. 
Himalayas (Kooloo, Nepal), Assam, Cacliai%Manipup.-7>^ qiqmitcus leuro- 

Tenasserira. — P/. iiiternicrlius. 
Mergui Archipelago. —It. hi/pomelcmus geminomm. 
Siani, Oambodja, Saigon.— Pi!, li/lei, hjpomelaiius condoreiisis. 
Pulo Condor. — Pt. hypumelanus coiidi/rensts. 
Formosa. — Pt. foniiosus. 
Soiilli Liu-kiu islands. —7^^. da-si/malliis, loochf/ensls. 

3. Indo-Malaya. 

Andauians. — Pt. iyileri, sati/rtcs, ? vampyrus. 

Nicobai-3.— P(f. melanoius, fammlus, ? vampyrus. 

Jfias. — Pt. niadicns. 

Eugano. — Pt. modiglianii , hyjMmelanus engamis. 

Christinas Island (S. of Java). — Pt. vaiulis. 

Borneo rind. Labuan, Mengalnn, Sibiitu, Lainboyan Islands). — /'/, oam- 

pyrii.f nututue. In/pomclanws tomesi, aprcioi'us. 
North Narunas (Bung.iran, Pulo Panjang. Palidak, Laut).-Pi'. vampm-us 

ncitiiiicB, hypoinchiius caimx. 
South Natuuas.— Pi*. hypo,iielanm annectens. (Pt. vampyrus naiunw, not yet 

Tainbelaii Islands, Pulo Anr, Tioman.— Pi", hypomclunus lepidm. 
Malay Peninsula. — Pt. vampyrus mcdacccnsU. 
8un'.atra (Linga Archiijelago, Banka).— P/. vampyrus malaccensis, h/pomc- 

unius tomesi. 
Java. — Pt. vampyrus vampynis. 
Bawean Island. —Pt. aferrimiis. 
Bali. — Pt. vampyrus plntun. 
Cagayan Sulii. — Pt. liypiimclanus cagaymius. 

Philippines. - Pt. vampyrm laiwnsU (generally), h/pomelanm cagai/anus 
(generally), specwsus (Malanipa I., off Zaniboanga), miiiius (Luzon) 


4. Anstro-Malaija. 

Celebes. — PI. Tiypomdaiius mfims.^aruus, mimus, dohsoni, akcfo, jwrsovntus 

Sanghir Islands.— P/". hypomelaims macassaricus, canicrps, chri'/suproctus 
melnnopogon. ' ' 

Talaul Islands. — PI. hypom^lamts macnssarirus. 

Bonerato, Dyanipea, Salayer. — Pt. gri.sciis, akcto. 

Lonibok. — Pt. vuvipyrus pluton, (d<xto, loudioccnsis. 

Suniba. — Pt. morio. 

Flores. — Pt. lombciccnsis. 

Alor. — Pt. ••ioUtnrius. 

Savu. — Pt. vampyrus cdulis, morio. 

Tmwe. — Pf. vampyrus cdulis, griscus, temmiiic/a. 

Gilolo group (Morotai, Gilolo,'Ternate. Batuhian). -P;". hypomchauisfiypi<me- 

lanus, caiiiceps, chrysa'ichen, pcrsonntus. 
Ghebi, Salawali, Mysol. — Pt. chrysaiwhcn, pajjuaitus. 
Sula Islands. — Pt. caiiiceps. 

Aiuboina group (Burn. Amboina, Ceram, Goram, Manavolka, Watubella). 

Pt. mclanoprigoii , vcularis, cJirysoprocfus, temmincki, persnnatus, ?arqeu- 
Banda Islands. — Pt. pallidus, mclaiiopogon. 
Timor Laut. — Pf. melauopogou. 
Key Islands. — Pf. Kryriisis. 
Am Islands.— Pi', arneasis, vmcrotis. [Cf. Pt. rubiginosns, fumigafn.', p. 213. 

86 riEEopus. 

New Guinea.— -P/. hypomelanus luteiis, chri/sauchen (N.W.), coiispicillatin 
(S.E.), papuanus, eptdarkis. 

Conflict, Trobi-iand, Woodlark, Kirhvina, Alcester Islamls.— T^, hypome- 
lanus Intens. conspieillafns, poliocephalus. 

Admiralty Islands. — Pt. admirali/atiim. 

Ei.»niai-ek Archipelago. — P^ neohiheriiicus, cnphtratns. 

Solomon Islands.— Bougainville, Shoi-tland : Ft. cul.oin's, grundis. Vella 
Lavella: Pt. lavellanus. Gliizo. Eubiana, New Georgia: Pt. solomoim, 
rubianiis, luoudfordi. Guadalcanal': Pi. reii/iie>-i, icoodfurdi. San 
Christoval : Pi. coynatug. 

5. Australia. 

N. and E. Australia.— P/. brniincuf, gouldi, cvinipirilMii^, poliocephalii^, 

6. rohjnesia. 

Benin and A'olcano Islands. — Pt. psdaphon. 

Mariannes. — Pt. marianiius, ? fiibcrfiiliifus. 

Pelew Islands. — P/. pcleweiisU, pilosus. 

Carolines.— Yap, Mackenzie: Pt. yapeiisiK Ruck A(oH: Pt. inmlaris. 

Murtlock: Pt. nolossiiius, phaoccpliaius. Ponapc : Pt. molossiinus. 

Ualan (Kusliai): Pt. i/alaiius. 
Sta. Cruz Islands (Vanikoro).— P/. vaiiil-orcntif, ? tnJicrailattis. 
New Caledonia.— P/. ornatiix, geddiei. {Cf. note on iV. retulns, infra p. 155.) 
Loyaltj- I.^lands. — Pt. aurutu^. 
New Hebrides {\^w\iewm).—Pf. geddiei, anctiauits. 
Vijis. — Pt. tonganus, nawaicnsis. 
Tongas. — Pt. tone/amis. 
Sanioas. — Vt. tcmgunun, ganwendg. 

Chronohgicnl list of species and szihsj^ecies. 

1605-1751. — The earliest known forms of Pteropus are the largest of the 
two species inhabiting Mauritius and Reunion {Pt. niger), the single Malagasy 
species {Pt. riifiis), the single Indian {Pi. gigaittnis), and the largest liido- 
Malavan species {Pt. vampynis). Pt. niger was described and figured by 
Clusius, in 1605, under the name Ves^pertUio ingens (Esot. libri decern, p. 94). 
Next in chronological order comes, apparently, Flacourt's ' Fany ' (Hist. 
Madag. p. 166, 1658), which is the Malagasy Pt. riifus ; Edwards's description 
and fio-ure of the ' Great Bat from Madagascar " (Nat. Hist. Birds, pt. ir. 
pi. 180), usually referred to as the earliest record of this species, is nearly a 
century later ( 1751). The Indian species {Pt.giganteus) was briefly mentioned 
bv the Swedish traveller Nils Matson (Koping) in his once ianious ' Reesa 
genom Asia,' 1667 ("Mycket stoora Nattblackor," p. 132), and soon found its 
wav to the European Curiosity Cabinets (Olearius, Goltorff. Kunst-Kanim. 
p. "24, pi. XV. fig. 1, 1674,' Vespcrtilio Indira'; JacobiEus, Mus. Reg. p. 12, 
1699. ' I'eipei-tiliones duo Indici'). Seba's Canis volans Tenmtcmus Orientalis, 
1734 (Thesaiir. i. pp. 91-92, pi. Ivii. figs. 1,2), is the large ludo-Malayan 
Pt. vampi/rns: the original of his fig. 2 is now in the collection of the British 
Museum. To these early records may perhaps be added Camel's ' Cabug vet 
Panicqui,' 1708, from the Philippines (Phil. Trans, xxv. p. 2198), which is 
not determinable with certainly, but must be either Pt. vampynis or Acerodon 

Brisson, Dauhenfan, Biiffon. — Two speries were known to Bri.sson, in 1756 
(Regn. Anim., based chiefly on the collections in the Cabinet Reaumur), from 
personal examination, both of them from the Mascarenes, viz. Pt. rufus aut 
niaer, 'La Rousselte,' which is Pt. niger, and Pt. f metis, anriciilis hrevibus 
acutiu!:ciilis, coJto .viperiore rvhro, the ' Roussette a col rouge,' which is Pt. sub- 
niqtr (h\s third species of " Pteropus." Pt. rnirirutis patitli^. is a Phyllostorae 
Bat, Vninpyrma speririim). Daubeuton's 'Chien-volaut,' 1759 (Mem. Acad. 

K. Sci. Pai'is, p. 384), is PL iiiger, his ' Eoiisseltp' probiiblj either ihe same 
species or I'i. »iif»iif/er. Buif'oii, 176o and 1771) (Hist. Nat. x. pis. xiv. and 
xrii., and Siippl. iii.), gives fisures and detailed descriptions of tho 'Koussette ' 
[PL 11 iqcr^ and 'Hou^eUe' [Pt. si/biiiqcr], together with an excellent account 
of the habits of tlie former species based on letters from de la Niix, wlio for 
more than M years was a resident of Keiinioii. 

Linne. — Seba's Cam's vulaiis TeiiutLaiius OrientaJis was bj Liune, in the 
second (1740) and seventh (1748) editions of tiie ' Systenia Naturte,' referred 
to as Vespertitio cauda nulla; in the tenth edition (1758) it was named 
Vesper/ilio vampi/ru^. in reference to its alleged blood-sucking habits. Tech- 
nically, the name PL vampi/riis must therefore stand for the large Tndo- 
Malayan specie.*, but other synonyms given by Linne are evidence that it was 
intended to cover any then known form of Fruit-bat (formally with the 
exclusion of PL siihiiif/er, inasmuch as Linne ignored Brisson's writings). 

1782. — The Indian species described and named, Vapertilio gigantea, 
Briinnich. — Number of technically named forms now 2. 

1792. — Mostof the early post-Linneiiu compilers recognized three '"' varieties " 
of P/cropiis (or Vcspertilh) vampyrui : a large [ = Pt. iiiycr + vampi/rus + rufus, 
or two, or one of these speci;s], a small \_ = PL subnigo-], and a straw-yellow 
[=Eidolun helvinn]. Kerr (1792) was the first to propose technical names for 
these " varieties," viz. J'esp. vampyrus iiigsr, siibniger, and heloits. — Number of 
named forms, 4. 

1797. — Ve^ertilio caniinis, Blumenbach [^J^'. vnmpi/rtin, L., 1758]. 
1802. — Spectrum ruhidum, Daudin [=Pt. suhniger, 1792]. 
1803.— E. Geoffroy's Cat. Mamm. Mus. Nation. d'Hist. Nat. (subseqnently 
suppressed by the author); descriptions of PLJ'uscu.s [= vigcr. 1792j, rufiis 
[earliest name of the Malagasy species], and ruber [= fubuk/er. 1792]. — Pt. 
fuscus, Desmarest, nee Geoffroy [the name 'Juscus' evidently an abbreviation 
of Brisson's diagnosis of the ' Eoussette a. col rouge,' see above; =^PL sub- 
iiiger^. — Named forms, .5. 

18t}4. — Hermann's Obs. Zool. ; Ve^p. cdano [ = Pt. vaiupgrus ram/n/rus, 
1758], iiudus [indeterminable, perhaps a young Ptcr('pus~\, and muuritiaium 
\^=niger, 1792]. 

1808. — PL rufus, Tiedemann, nee Geoffroy [the name ' 7-ufus ' an abbrevia- 
tion of Brisson's diagnosis of ' La Eoussette,' see above ; =PL iiiger. 1792]. 

1810. — E. Geoffroy's classical paper in Ann. Mus. d'Hist. Nat. xv. ; five 
forms described, two of which were new : PL edulis [yampyrus edidU]. from 
Timor, edwnrdsi [ = n(fus, 1803], vulgaris [=uigcr, 1792], rubricollis [ — sub- 
niger, 1792], and qriscus, from Timor. — Named forms, 7. 

1814, 1815, 181(j.— 7V. torquatus, G. Fischer [=suh),iger, 1792]. Pt. cdlaris, 
Illiger \^ = siilmigerl^. PL madaqascariensis, Oken [ = /-z//'«,s, 1803]. 

1820, 1822. — Desmarest's Mammalogie ; I'L junr/iicus [^vamjjyrus vofti- 
pyrus, 1758], mariaiiinis (Guam, Mariannes). — Named forms, 8. 

1824. —Voyage of the ' Uranie'; Pt, keraudren, Quoy & Gaimard [ = inari- 
aiiniis, 1822]. 

182.5. — Teniminck's Alonographies de Mammalogie, vol. i. ; six supposed 
new forms, four of wjiich were really undescribed: Pi. nieditis [—giganfeus, 
1782], phaivps [ = rufus, 1803], policcephalus (Rrst Australian species known), 
dasymallus ("Japan," really Liu-kiu Islands), pallidus (Uanda Islands), and 
personatus (Ternate). — Named forms, 12. 

1828, 1829. — Pi. du.^i(mieri, Is. Geoff, [indeterminable, externally like 
PL vanikorcusisl. Pt. pscla2}hon, Tradescant Lay, from the Bonin Islands. — 
Named forms, 13. 

IS^X — Voyage of the 'Astrolabe'; Quoy & Gaimard: Pt. vaiiikurciisis, 
Vanikoro, and toiigaiius, Tonga Tabu. — Named forms, 1.5. 

1835.— Hodgson : Pt. leucocejthatus, Nepal [gigaiiteits kucocrphalus]. — 
Named forms, 16. 

1837. — Tlie second volume of Tennninck's Men. Mamm. : Pt. fuiiereux 
[—rampi/rus edulis, 1810], chrysopructus, Amboiua, and aleclo. Celebes. — 
Named forms. IS. 

1839.— iV. (issa>iicusif, McClelland [—gigaiiicus leucoccplmlus, 1835]. 

1842. — Hoiiibron and Jncquinot (V'oy. Pule tiLul) ; Ft. irifularis, Caroline 
Islands. — Niinied roriiis, I'J. 

1844. — Gray (Voy. of tliB ' Sulphur ') : I'l .a r<jeiitutU6,pYe%\i\n\\\)\^- Aiiiboina. — 
Kamed forms, 20. 

1848.— Peale (U.S. Exploring Expedition) : Pt. vociferus, Mangsi I., Balabac 
Straits (indeterminaljle, perhaps Pt. ki/pojiielanus), and mmuensis. — Named 
forms, 21. 

1850. — Pt. coJispic Hiatus, Gould (second Aifcttralian species; cf. 1825), — 
Named forms, 22. 

1853. — Teuiuiiiick's E--q,iiisses Zoohigiquea : iV. pluton, Bali and Lonibuk 
[vampyruti plutijti], /(■iicopteniii, Fhiiipyhu'S, h//pomc(auiis, Ternate, and molus- 
i<iiiiis, unknown locality [Caroline.'*]. — 'Named forms, 2(). 

Ig58. — Gray, on Wallace's Aru Islands collections : Pt. argeniatns [name 
preoccupied, 1844: species redescribed by Peters, 1867, as Pt. melanopoyuit 
var. antensis], 

18tiO. — Pt. qcddiei, MacGiUivray, New Hebrides.- -Named forms, 27. 
1862. — By Peters : Pt. scapulalus (third Australian species known, cf. 1825 
and 1850), and chi-i/saachcn, Batehian. — Named forms, 29. 

1863, 1866. — Pt. i'eA«^/fs, Jouan, New Caledonia [</. p. 155]. Pt. ladanotus, 
Blyth, Nicobars. Pt. Uvinystonci, Gray, Comoros, — Named forms, 31. 

I8ii7. — Peters's revision of the genns : Pt, ocularis, Ceram, macrotis, " Bnrn " 
[really Aru Is.], melaiiopogon, Amboina group, melanopogon var. aruensis, Aru 
Is. [iV. aracii>ii»], inclancpiH/dit var. /aj/eiixis, Key Is. [Pt. /cegciis/s], tenimhicki, 
Ambouia, and (joiiltii (fourth yliustralian species desei-ibed, cf. 1825, 1850, 
1862). — Rosenberg's Reisnaar de Zuidoostereilauden ; Pt.ritliif/uiosus [^?aru- 
ciisi,s\,fui)Hgatu!i i—?aritejisi8'],a,nd liisignis \=macnitis\. — Named forms, 38. 

1868.— Peters, on the Marquis Doria's Bornean collections : Pt. hyponicla.ims 
var. tomcsi. — Named forms, 39. 

1869. — Zelebor (-Novara' Expedition): Pt. iiicobaricui< [=inelaHotus, 1863]. 
Petei-s, revision of nuiterial in the Paris Museum : Pt. condorensis, Pivlo 
Condor [h//po7neknitis condi>rensis], a.nd tuhcrctilatus, unknown locality [probably 
either Vaiiikoro or Guam]. — Named forms, 41. 

1S70.— Gray, italogue of Monkeys. Iiemurs, and Fruit-eating Bats; sixteen 
supposed (six' really) new forms: Spectrum anetinnum, New Hebrides, Pf. 
kclaarti [=gUjaiUeus g/gautms, 1782], mysoleiisis [=chrgsaucheH, 1862], mggol- 
eiisis var. ceramcnsia {—ocularis, 1867], oruatus, New Caledonia, looclwiims, 
Liu-kiu Islands, caviceps, Batehian, uawaicnsis, Eijis, JlavicoHis [ = tonganu,'>, 
1830], [=ca)iiceps], tricolor [=:hi/poimlai!Us hypomelanus, 1853], raijncri, 
Guadalcanar, cUeyi \_— scapulalus, 1862], vitiensis [=/iawaic)isis], mcwkluti var. 
batchiana \_ = caniceps\ and chincnsis [^leiwopterus, 1853]. — Named forms, 47. 

1873, 1874.— P/,, Formosa, P. L. Sclater, Pt. whitmeci, Alston 
[=sam(A'i/di, 1848].— Named forms, 48. 

1876. — Peters (' GazwUe ' Expedition) : Pt. capistratus and melanopogon var. 
neohihernicus [I't. ■ncoJdhernicmi], both from the Bismarck Archipelago. Same 
author (dealer's specimen) : Pt. dcyencr, stated to be from the Aiu Islands, in 
reality from the Bismarck Archipelago [-^Pt .neohihernicus]. — Named forms, 50. 
1878. — Dobson's Cat. Chir. B. M. : Pt. Imnbocensis, Lombok, rodricci/sis, 
Rodriguez, hninncus (fifth and last Australian species described, cf. 1825, 1850, 
1862, 1867), pteronotus \_=vampyrus vampyrus, 1758], and fuscits, Celebes 
[name preoccupied ; renamed, 1908, Pt. dobsoni, K. And.]. — Pt. epulariiis, 
Ramsay, New Guinea. — Nan\ed forms, ^4. 

187y__13obson, on material from the Paris Museum : Pt. germcdni, New 
Cidedonia [type ajiixirently lost ; = vetuliis, 1863]. 

1881.— Peters and Doria, on Dr. O. Beccari's collections: Pt. melanopogon 
var. papnana. New Guinea [Pt . papiiamis]. — Named forms, 65. 

1882. — Thomas, on specimens from the Godeffi-oy Museum : Pt. pkegocrjjhaliis 
and brevic.eps, both from the Carolines [the latter withdrawn by Thomas, 1887 ; 
=molossinus, 1853]. — Named forms, 56. 

1883.— Peters (Dr. O. Finsch's Caroline Island.-; collections): Pt. ualaniis ; 
]'t. iiiidf^dnns. bised (1853) on specimen 'Vom uulniunn lucalily, rediscovered 
in P')niipe.— Named forms, 57. 



f j^f^'''-7,%/-'-''"'"'"s : Pf.gramU,. Shorthuul, Solomon Islands (C M VVood- 

1<^8.-By Ihomas: Pt. woodfonU, Guadalcanar, Solomon Island, and 
covmmim{ = mohiheniicus, 1876].-Nanied forn.a, 61 ' "'^ 

[-S/«4 l?iT 'T:^'^''"- ^l'™. "««>""- Isl^^nds," in reality Carolines 

ThlnuS-^ p*/'::'^r'/' -^•'■' i^^°"'' collections : Pf. aldchrensls, Aldabra. By 
iirled Ibrm,, ef ^ '""' ^"°'''"°' '*'''* «^"">«'^-'«'««> Adn.iralty Islands.- 

6ar/r«sJ r//ri.««,Y/yr«., l = ke,,ei,s,,, 1867 , and /^vW/ZoHnauie nreoccuDied ]87()- 

i'ku i';r-'''"^^° (%^'""«^«"«.s ^««»m..««,].-Na™ed forms, gg/'"""""'""' 

Mearns ir^.^ J Prr^ ■^'^- f^^""'''"'^^' Central Solomon Islands. By 
JMeanis . It. laiiams, Pliilippnies [vami»jms lanensisX and caaavanu^ C-^J 
^'"^".f"'" [Aw^"'f'f«««.sc«ya^„;,.,.]._Nan.ed forms, Tl. '"^"y"""'^ C!aga- 

lJU6.--ByM.ler: PA havcauus l^alen-imus, 18091 ,«-«<i/«,s Nias and 
"'^^^n' ,^:',f "« %^'""*'"« ^"<7«««.-].-Named forms, 74. ' ' """^ 

Oct 1908, Feb. 1909.-K. Andersen, revision of genus : Pt.h»pomela>ms canu^ 


../^^«,-/«.Alor Island, rufus pr.iccps, S. M^^^^^'^^l^a^S^ Sr' 
«.«^.«,s, lenassernn, .«..,,,/.„.. ,ncUaccen.i., Malay P^nh, la T .2;,/^ ]^" 
Natunas, ;«or(o, Siiniba,y.ite«.s-, Pelew Islands r^/K, ;;/ P.iJk "«^«"«. /I- 

Loyalty Islands.-Nan.«l form; 99 ' ' ^'''^''' '"'' '^'"■"^"*' 

Two-thirds of the 99 known forms have been described bv six anthor. 

Tlie types of the 99 forms are distributed as follows :— 
fngentatu,, camdns, hrtmneus, caniceps, camis {hypomdamis c\ Z.aln: 

gham, (,;otype in Genoa Museun,), mono, natalis%atu2\TaZ;uTi\ 

N™n( 1) Sc?a,?r ?''?h^'°'M^^; Y^ (1). Linne (1), MacGillivray 1 
4, V^' *'^"'''^*^ (^)' J-nomas (7;, Andersen (2'') • ^ '^ 

Lejden Museum 11: Pi. a/ccto, chrg.aproetus, d<u<,,mallm, hvpomelanv. 
fo^Zf •''''ir'r''''AV'''"'"^''''"*' (""'-^P^ '-^ ^«"* Museum,, molosJ'^Sa^ 


U.S. Niitional Museum, 9 : Pt. aldahrensis, cagrtyaniis {hypomelairus c), 
enganus {hijpnmclnnus e.), fmumhia, geminorum {hypomelamis g.), lanensis 
(rampijrics I.), lepidiis {hi/pomclanus I.), niadicus, samo'ensis ; described by 
Mearns (2), Miller (5), Peale(l), True (1); 

Paris Museum, 8 : Ft. edulis (vampj/nts c), grUeus, insularis, mariatmus 
(cotyjie in Leyden Masenm), segckelleits/s, tonganus, tuberculatus, vaniJco-rensis ; 
described by Desmarest (1), E. Geoffroy ^2), Hombrcn & Jacquinot (1), 
Milne-Edwards (1), Peters (1), Quoy & Gaimard (2) ; 

Genoa Museum, 2: Pt. modiglianil (cotj'pe in British M-xxsenm), papiianus ; 
described by Peters & Doria, and Thomas ; 

and one each in the Oopenliagen (Pf. giganfeiis, Briinnich), Calcutta (P/. 
melaiioUis, Blyth), ?Zi-ka-wei (Pf. h7jpomdanm macassarictis, Heude), Sydney 
(P(. epulariits, Ramsay), and Cambridge (Massachusetts) Museums {Pt. ariel. 
G. M. Allen) ; one is presumably in private possession {Pt. ti/tleri) ; while fi^e 
are apparently not in existence [P/. niger. poliocephaliis, rufag, subitigcr, toniesi 
{hypomelamis t.)\. 

Synopsis of Groups and Species *. 

A. Posterior basal ledges of large premolars 
I. >Skull and dentition unmodified Pteropine f. 
Tibia generally naked above J. 
a. Dentition not unusually heavy. Size 
moderate or.«mall: forearm 94-171 mm. 
a'. Interfemoral scarcely developed in centre, 
ft'-. Ears moderate or short, nt)t narrowly 
u\ Colour varying, but never blackish 
above and beneath with sharply con- 
trasting light yellowish mantle .... A. Pt. hypomelamu.s 
«'. Cheek-teeth of normal size: breadth group, p. 98. 

of p' about one- third that of palate 
between p*-p*. 
a'. Tibia naked above. 
«". Skull larger : total length 61 --'j- 
68v mm. 
a'. Fur shorter : 10-14 ram. on 
back. Forearm 121-146 inni. 
(Mergui Archip. to New [p. 101. 

Guinea) 1. Pf. hypoiudunus, 

b~. Fur longer: 18-19 ram. on bark. 
(Blackish above and beneath 
with pale mantle and centre of 
breast.) Forearm 1;39 mm. 
(Andamans) 6. Pt. satynts, p. 142. 

* The geographical review of the species and subspecies (pp. 81-8(5) 
and the list of the species arranged according to the lengths of forearms 
(pp. 97-98) will often greatly facilitate the identification of the specimens to 
be determined. 

t Except the Mascarene Pf. suhniger, in which the cheek-teeth are small 
and unusually narrow, and the skull modified accordingly. 

* The tibia is naked above in all species of this secli(ni, except in seven 
peripheral species of the Pf. hypomelanus group, viz., Pf. solomoiiis (Central 
Solomon Is.), firunneiis (E. Australia), ornatus (New Caledonia), ai/rafns 
(Loyalty Is.), drtiiymof/iif: (Liu-Kius), forii'osits (Formosa), and ■•'iifii//'/cr (Mas- 
careues), and in the single Mascarene species of the Pt. ni/iis group (Pi. iiigcr). 


6". Skull siuiillfr; tulal length oi'o- 
oiJ'S mm. 
c^. Ears from orifice 2o-26 mm. 
ft''. Forearm 120'5-12.'> mm. 
«■'. Darker : underparts tawny. 

(Sibutii I. ; Malanipal.) . . ." . 2. Ft. xpetiosits, p. 132. 
6". Lighter ; underparts golden 
ochraceous. (Philippine.*; Ce- 

lebe.i) 3. iV. inimus, p. lo.3. 

l/^ Forearm 113-119 mm. 
t''. Flanks dark brown. (Canda 

Is.) 4. PL j)o//idus, p. ]y(3. 

(P. Flanks pale like underparts. 
(Timor; JJuncrato; TJyampea : 

Salayer) .">. PL (jri^em, p. 137. 

(P . Ears from orifice 21-22 mm. 
c''. Forearm 11 8-12(j mm. 
t^. Underparts ochraceous - buf¥y 
tinged with cinnamon. (Nico- 

bars) 7. Pt.faunubis, p. 143. 

y*. Underparts seal-brown heavily 

nii.xed with c-reyish. (Ad- [p. 144. 

miralty Is.) B. PL (ichiiircilifatum, 

(7\ Forearm 109-114 mm. (Un- 
derparts dark brownish.) (W. 

Solomon Is.) 0. PL colonus, p. 147. 

A\ Tibia clothed above, at least for 
proximal half. 
c'v Fur of back directed posteriorly. 
('". Forearm about 110-120 nun. 
Mantle russet, cinnamon, or 
e". c-m" * less than 22 mm. Fore- 
arm 110 mm. (Central Solo- [p. 148. 

mon Is.) 10. PL solomonis, 

f\ c-m- more than 22 mm. Fore- [p. 149. 

arm 118 mm. (E. Australia). 11. Pt. bnameus, 
f. Forearm about 145- lo2 mm. 
Mantle bufly. 
(f. Head, back, and underparts 

dark brown. (2sew Caledonia) 12. 7Y. (;;•«'?;(/.<. p. l.")3. 
h". Head and underparts ochraceous, 
back mixed ochraceous and 

brown. (Loyalty Is.) 13. /')'. rtM/Y(^«s, p. loti, 

6^. Fur of back long and spreading. 
<f. \'aried with bufly on back. Fore- 
arm 125-137 mm. (S. Liu-kiu p. 1.59. 

Is.) 14. PL dcmjma'Uus, 

h~ . Xearly uniform dark brownish 

above and beneath. Forearm [p. ]()3. 

130-137 mm. (Formosa) .... 15. Pt.fvnnosiig, 
6*. Cheek-teeth reduced : breadth of p' 
le.-:s than one-fourth that of palate 
between ji'-p'. Eiiri very small, 
hidden in fur : tibia thickly clothed. 
Forearm 95-90 mm. (Mascarenes) Iti. PL subiv'gcr,\^.l(^i:. 

* Maxillary loolh-row, from front of eunine to back of last molar. 

S2 rxEiiorrs. 

U\ Blackish above and beneatli, often 
sprinkled with pale gTeyish, and with 
sharply contrasting- light yellowish 

mantle . U. Pt. mariaxnus 

c'. Small: forearm about 113 mm. guoup, p. 172. 

(Pelew Ts.) 17 . Ft. iielewensis, 

(l\ Larger; forearm 13J-lo4 mm. [p. 173. 

c'\ Eyes smaller: orbit ll-o-ll'8 mm. 
c*. Fur shorter: 9-11 mm. on back. 
f. Dentition weak ; length of p^ 
8-8-4 mm. Forearm loO- 

134 mm. (E. Carolines) 19. Pt. na/n/ui.s-, p. 177. 

j'. Dentition stronger : length of p ' 
4-5—5 ram. 
i^. Dentition moderate : length of 
p^ 3-9-4'l mm. Forearm about 

1."10 mm. (W. Carolines) ... . 18. Pf.>/(ij>c)isi-',\). 17^. 
_f. Dentition strongest : length of 

p-i 4-8-0 mm. Forearm 134- [p. 178. 

137 mm. (Marianne Is.) .... 20. Ft. mariannus, 
p. Fur longer: 14-17 mm. on back. 

Forearm 135-143 mm. (S. Liu- [p. 181. 

kiu Is.) 21. Pt. loocho'ensi», 

(P. Eyes larger: orbit 12-5-13-8 mm. 
(/''. Smaller : c-m'-^ 24-5-2(), forearm 
136-150 mm. 
Ic'. Shorter-winged: third metacarpal 

about 88-89, iorearm 136- [p. 184. 

137mm. (Sta. Cruz Is.) 22. Pi . ranikm-i'nsis, 

P. Longer-winged ; third metacarpal 
about 93-5-102-5, forearm 139- 
150 mm. (Fiji ; Tonga; Sa- [p. 186. 

moa) 23. P/. tviujanus, 

h^. Larger : c-m- 27'8-30, forearm 
about 154 mm. (New Hebrides ; 

New Caledonia) 21. Pt. /jedJUi, ]\.\S^i). 

h\ Ears longer (reaching back of eye), 
narrowly pointed above *. Dentition 

heavier C. Pt. cAxuiavs ouour, 

c^. Ears about 24 mm. (orifice to tip). [p. 192. 

Back sepia suffused with golden 

buffy. Forearm 145 mm. (Celebes) 25. Pt. (J'jbsoiil, p. 192. 
rf\ Ears about 31 mm. (orifice to tip). 
•j t'^. Dentition stronger: length of m, 

6" 1-5 "5 mm. : fur short. Forearm 
135-140 mm. (Gilolo group ; 

Sula Is. : Sanghir) 26. Pt. rantcep'^, p. 194. 

/'. Dentition weaker; length of nii 

about 4-7 nun.; fur longer. [p. 197. 

Forearm ? 136 ram. (? Amboina). 27. Pt. aiujcutntus, 

b' . Interfemoral distinct in centre. 
C-. Ears strongly attenuated above, sub- [p. 200. 

acutely point -d. . D. Ft. rufus ftnour, 

* Exempt Pt. rtuhsoiii, ill which the ears are nearly similar in form to tliose of 
I'i. hijpoiii'ji' 

rrEitoi'us. 93 

e\ Tibia naked above. Ears exposed. 

No colour contrast between lateral 

and median tract of back. 

g*. Kars much longfer : 37-38 mm. Size 

larger : forearm 158-171 mm. 

(Madagascar) 28. I't. rafus, p. 20-> 

h\ Ears much shorter: 31-32 mm. 

Size smaller: forearm 13-1-157 mm. 

«''. Hack blackish, uniform or sprinkled 

with gieyish. Larger: forearm 

143-157 mm. 

i'\ Back not, or extremely thinly, 

sprinkled with greyish ; breast 

and belly yellowish buff. Forearm [p. 208. 

151-157 mm. (Comoros) 29. I't. comorcnsis,' 

,/''. Back consjiicuonsly sprinlvled with 
greyish ; breast and belly much 

darkened with brownish." Fore- Tp 9\o 

arm 143-154 mm. (Seychelles). 30. Pt. ^ei/chellen'sfs,'" 
/'. Back strongly tinged with broccoli- 
brown. Small ; forearm about [p. 213 

, ,^.,.l^-*,'"™- (Aldabral.) •■i\ . Pt . aldabrens'is, 

/ . libia clothed above. Ears cearlv con- 
cealed in fur. Buffy sides ot" back 
contrasting with dark spinal tract. 

Forearm 159-171 mm. (Mascarenes) 32. Pt. nvjer, p. 215. 
d-. Ears rather broadly rounded off at tip . E. Pt. melanotus 
(f. Centre of breast light-coloured. group, p. 223. 

i^. Back blackish seal-brown. Forearm Vp' 224' 

153-165 mm. (Nicobars) ^Z. Pt. melanotics' 

J . Back approaching hair-brown. Fore- 

, , ^. ^l^ 153-ltJO mm. (Xias ) 2,0. Pt. niadicus, p. 229, 

A*. L nderparts uniform blackish. 
/.:'. Forearm about 150 mm. Mantle 
sometimes pale-coloured. (Anda- 

„ ^,»ians) ':iL Pt. tytlevi, p. 227. 

/'. I'orearml2o-141mm. Mantle never 

(f. Skull (51-5-67 ; fur of back 18 ; fore- [p •7o-> 

,- «™ 13-1-1-11 mm. (Engano) .. ZQ. Pt . modiulianiL "' 
h\ Skull 04-5-56 ; fur of back 25 ; foie- 

arm 12.3-135 mm. (Christmas I.) 37. Pt. natalis, p. 233. 
ti. Dentition unusually heavy. Large : fore- 

ai-m l(i2-204 mm ! p. Pt. mklaxopogox 

c._ Lars not peculiar. Forearm 179 204 mm. ghoup, p. 237. 

e-. Furred area of back restricted to 
narrow spinal line. Forearm 196- 
204 mm. (Amboina group; Banda; [p •:>38 

Timor Laut) .-8. Pt. mdunopoqmi, 

/-. F urred area of back of normal breadth, 
j'. Back silvery whitish. F'orearm about [p 241 

190 mm. (Aru Is.) m. Pt. avuemis, ' 

J . Nearly uniform yellowish above and 
beneath. Forearm 179-188 mm. 

(Key Is.) 40. 7V. /;,.>/ensis, p. 246. 

d. Lars semicircularly rounded off above. [p oj; 

Forearm 162 172 mm. (Comoros) .. 4\. P/. lianpstmiei.'' 



II. nostrum sliorteueil, deutition more or les« 
modified. Tibia nearly always clothed 
above *. 
c. Dentitiou heavy. Fur never silvery 
whitish or bufly above and beneath. 
e'. vn^ always, m- often, conspicuously re- 
duced ; u and p, normal. 
(/-. -Hostrum less shortened ; i, not reduced, 


k'. Tibia densely furred above, at least 
for proximal half. (Solomon Is.). 
'/«'. Brownish above and beneath with 
cinnamon or russet mantle. Fore- 
arm at least 121 mm. (San Christ- 

n-i. Upperside tri-coloured : briyht 
mantle, dark back, bright rump. 
t! *^. Smaller : forearm 137-141 mm. 


j\ Ijarger: forearm 151-172 mm. 
i\[ h'^'. Crown and face grizzled with huffy 

and greyish. 
w(". Lower leg a]x)ut7(i"o mm. ; colour 
much lighter. Forearm about 

I(j3 mm. (Rubiaua) 

n'. Lower leg about 6;)-68 mm.; 
colour much darker. Forearm 
151-156mm. (VellaLavella). . 
,." P. Crown and face blackit^h seal- 

brown. Forearm 167-172 mm. 
(^Bougainville; Shortland) .... 

,'"' P. Tibia thinly haired above. Upperside 
generally tri-culourtd. Forearm lOy- 

177 mm. (Amlxiina group) 

/(-. Rostrum much shortened; ii reduced, 


«(•*. Tibia furred above ; forearm 108- 

127 mm. 
v*. Fars exposed. 
k^. Larger, darker. Foreaim 113- 
122 mm. (Lombok ; Flores) .... 
fi. Smaller, paler. Forearm about 

108 mm. (Alor L) 

p*. Ears nearly concealed in fur. Fore- 
arm 124-127 mm. (Rodriguez) . . 
H-K Tibia nalced above. Very small : 
foiearm 94-99 mm. (Carolines) . . 
/'. ni;,, m", and ii normal ; i; and pi en- 
larged ; rostrum very short. 
i^. Posterior ledges of upper incisors of 

normal breadth 

o^. No inner basal ledge in lower cheek- 


[p. 250. 

42. Pt. ccMjnatus, p. 251 . 

43. Pt. rayneri, p. 253. 

44. Pt. n/tiianifs, p. 255. 

[p. 258. 

45. Pt. hivellanus, 

40. Pt. (jrandk, p. 259. 

[p. 2G0. 
47. Pf. chrystqiroctiis, 


Gitoup, p. 265. 

i"p. 266. 

48. Pi. loniboccnsis, 

[p. 269. 

49. Pt. solitarins, 

[p. 273. 
iX). Pt. ruilncensts, 

; p. 275. 
51. Pt. mohi^sinus, 


[p. 280. 

* The tibia is clothed, and generallj' thictlj clothed, above in all species of 

this .seetion except Pt. /i(olossiiiii.% iiisii/arif!, and pho'ticeplwlus (Caroline Is.). 


q\ i^ and Pi .■^lifilitly eiilai'ged. Fortsinu 

124-132 mm. " (Fijis) 

r-K \., aud p, much enlarged. Forearm 

about 144 mm. (.Samoas) 

;j\ A broad inu.'r basal ledge in p^, m,, 
and m,. Forearm 123-13U mm. 
(New llebride.s) 

y^. Posterior basal ledges of upper incisors 

unusually broad 

q"^. Tibia naKed a])ove. Small : fore- 
arm 101-109 mm. (Carolines), 
.s-^. Back dark brown. (Ruck atoll) 
t*. Back golden cream-bull'. (Mort- 


r3. Tibia furred above. Larger: forearm 
120-1.51 mm. 
u*. Upper molariform teeth not shortened. 
III". I pper canine without secondary 
cusp. Larger : forearm 132- 
151 mm. 
m''. Feet hairy above ; fur of back 
30 mm. Forearm 132-141 mm. 

(Bonin and Volcano Is.) 

n'^. Feet naked above ; fur of back 
20 mm. Forearm about 151 mm, 

(Pelew Is.) 

n'. Upper canine with small secondary 

cusp. Small: forearm about 120 

mm. (? Marianne Is. ; ? Vanikoro) 

r*. Upper molariform teeth shortened, 

subquadrate. Forearm about 140 

mm. (Philippines) 

<1. Dentition weak or even degenerate. Un- 
usually pale-coloured: silvery whitish 
or silvery buffy above and beneath .... 
y' . Head not striped. Forearm 94-101 mm. 

(Amboina group ; Timor) 

h'. Head striped. 
Ic^. Cheek-teeth of normal breadth. Fore- 
arm 110-116 mm. (]3ismarck Archip.) 
I'. Cheek-teeth excessively narrow. Fore- 
arm 86-96 mm. (Gilolo group ; Ce- 

B. I'osterior basal ledges of premolars prac- 
tically obliterated. (Tibia naked above in 
all species, except Ft. poliocephahis.) 
e. Cheek-teeth of normal breadth (breadth 
of p' 3, rarelv :f, that of palate between 
i' . p3 and p'' normal. 
}«-. Furred area of back of normal breadth 
(not restricted to narrow spinal line). 
Middle divided palate-ridges less 
than 8. 
«^ Ears long and sharply pointed 

[p. 280. 
52. Ft. nmcaiennis, 

[p. 284. 
63. Ft. samoensis, 

[p. 288. 
51. Ft. anetianus, 

J. Pt. pselaphon 

GROUP, p. 293. 

55. Ft. i/isularis, p. 295. 

[p. 298. 

56. Ft. pha-occjihalus, 

[p. SOL 

57. Ft. psefaphoii, 

58. Ft. pilosu.% p. 300. 

[p. 309. 

59. H. tnberculntus, 

[p. 311. 

60. Ft. leucoptems, 

K. Pt. temmincki 

GROUP, p. 315. 

61. Pt. temmincki, 

[p. 316. 
[p. 319. 

62. Ft. rapmtiutiis, 

[p. 321. 

63. Ft. persoiKitits, 

L. Pt. v.^MPYRrs 

GHOip. p. 324. 


?(;'. Smaller: foveanu 148 -lu4 mm. 

(Siam ; 8aigou ) 

.1'^. Lai'frer ; forearm lGO-220 mm. 
o\ Underside of body paler than back. 
o''\ Itatber larger, with slendei-er ros- 
trum and smaller teeth. Forearm 
163-177 nmi. (India; Ceylon; 


p^. Rather smaller, with broader ros- 
trum and heavier teeth. Forearm 
about 160 mm. (Maldives) .... 
2^. Underside of body quite, or nearly, 
as dark as back. 
g''. oinaller^ foreneck much lijjhter 
than i)reast. Forearm about 

180 mm. (Tenasserim) 

»•'•. I^arger; foreneck subsimilar to 
breast. Forearm 18.3-2i'0 mm. 
( Malacca ; ludo-Malaya ; Lesser 

Sunda Is.) 

P. Ears not low^ and sharply pointed. 
(/'. Blackish above and beneath, but 
never with light yellowish mantle. . 
q\ Smaller: forearm about 141 mm. 

(Sumba ; Savu) 

)■'. Larger : forearm 152-176 mm. 
i«. Orbit smaller: 13-7- 13-9 mm.; 
dentition heavier: length of p^ 
4"6-5*l mm. Forearm 1.52- 

160 mm. (Bawean I. ) 

f. Orbit larger : 14-2-14-8 mm. ; den- 
tition weaker; length of p' 4-1- 
4-7 mm. 
.o~. Cheek-teeth broader ; breadth of 
p^ 'i of length. Forearm 160- 
175 mm. (Celebes ; Salayer ; 


jr. Cheek-teeth narrower; breadth 

of p' J of length. Forearm 

153-180 mm. (Australia) .... 

z\ Blackish above and beneath with 

sharply defined light yellowish 


ji'\ Larger: forearm about 157-181 mm. 
ir. No well-defined spectacles. Fore- 
arm about 175-183 mm. (Gilolo 

group ; \V. New Guinea) 

r''\ Circumocular space and lores pale- 
coloured. Forearm 157-181 mm. 
(E. New Guinea; Australia) .. 
t\ Smaller ; forearm about 135 mm. 

(Amboina group) 

)fi. Furred area of back (in adults) restricted 
to narrow spinal line. Palate-ridges 


M^ I^iickdark vandyck-brown. Forearm 
about 190 mm.' (New Guinea) .... 

66. I'f. lylpi, p. .339. 

[p. 326, 
64. PL f/ignnfeus, 

65. Pt. ariel, p. 335. 

6/ . Pt. i/iteniiediu.^, 

[p. 343. 
68. Pt. vampyrus, 


71. Pt. mono. p. 370. 

69. I't.atennnus, 

70. Pi. aleclo, p. 365. 
72. Pt. (jouldi, p. 370. 

N. Pt. coxsriciLLATrs 
UKoup, p. 375. 

[p. 375. 
7.3. Pt. chrysauchen, 

[p. 378. 

74. Pt. con.'ijjictllatns, 

75. lU. aculari.s,!^. 381. 

[■GEOUP, p. .384. 

0. Pt. neoiiii!kenicu.s 

[p. 385. 

76. I't. pajjiuitiw, 


f-1. Back lifjliter, from biifty to 

iiuirs-brown. Forearm 187-199 mm. [p. 387. 

(Bismarck Arcliip.) 77. Pt. mohihernicus, 

j'. p' and p' sulj^quarisli. Ears long and 

.sharply ])oinled P. Pt. macrotis group, 

«". I'ibia naked above. (Blackish above [p. 392. 

and beneatli with sharply contrasting 
light yellowish mantle.) ' 

n-'. Larger: forearm 136-141 mm. (New [p. 392. 

Guinea) 78. Pt. epularius, 

.v^. Smaller : forearm about 120 mm. 

(Aru Is.) 79. Pt. maa-otis, p. 39U. 

]>-. Tibia thickly clothed above. Fore- 
arm about 160 mm. (Australia^.... 80. Pi. poliocephalus, 
f. Cheek-teeth excessively narrow (breadth [p. 397. 

of p< 4-i that of palate between p^-p^) . . Q. Pt. .scapulatus 
li' . Ears long and i)ointed. -Much larger: group, p. 402. 

forearm 131-143 mm. (Australia).,.. 81., 
V . Ears small, nearly concealed iu fur. [p. 403. 

Alueh smaller : forearm 92-99 mm. 

(Solomon Is.) 82. i'^. tvoodfordi, 

[p. 407. 
Lengths of forearms of species and subspecies * . 

8fi-80 miTi. : — Pf. persnnaftcs (Celebes ; Gilolo and Amboina groups). 

90-99 nrn. : — Vt. molossinus (Carolines), pnrsonatus (Calebes ; Gilolo and Am- 
boina groups), .•iiih/iitfer (Mascarenes), teinminc/d (A;nboina group ; 
Timor), woodfordi (Solomon Is.). 

100-100 mm. : — Pf. colonuh (Sbortland, Solomon Is.), intularis (Ruck group, 
C.irolines), pitaorcpkauis (Mortlock group, Caroliue.s), so/itarius (Alor 
I.), solmnoiiis (Ghizo, Solouiou Ib.), tonminc/ci (Amboina group ■ 

110-119 mm.: — Pi. admiraUtafiim (Admiralty Is.), hrumieus (Austi-alia), 
rapist rat lis (Bismarck Arch.), cnlcmus (Sbortland. Solomon 1a.). faunulus 
(Nicobars), ffrisens (Timor, Bonerato, Dyampea), lomhocensi^ (Loiubok; 
Flores), pdlidiis (B;uida li.), peleweiisis (Pelew Is.), solomoim (Ghizo, 
Solomon Is.), tuherculafus (? Sta. Cruz Is. ; ? Mariannes). 

120- 129 mm. : — Pf. admirnlifafum (.■Vduiiralty Is.), anefianKs (New Hebrides), 
corjnaius (San Cliristoval, Solomon Is.), dasi/mallus {lAu-kwe.), fan nidus 
(Nicobars), hypomelnnus (Indo- and Austro- Malaya), lombocensis (Lom- 
bok; Flores), »)^''ro//.s (Aru I.S.), w}/«7« (Celebes ; Philippines), natalis 
(Christnia.' I., S. of Java), nnviaicnsis (Fijis), rodricensis (Rodriguez), 
speciosus (Siiluls.), tuhcrcidafus (? Sta. Cruz Is. ; ? Mariannes), yapcnsis 
(W. Carolines). 
130-139 mm. : — Pf. aldahrensis (Aldabra), aaetiamis (New Hebrides), 
'{ argentatus (Amboina?), caxw^ps (Gilolo group ; Sula Is.; Sanghir), 

* When two or more species of Pferopus occur together in one place, thev 
are generally conspicuously different in size. The length of the forearm of a 
specimen to be determined, together with the precise locality given on its label, 
will therefore in most cases help the non-specialist to an easier and quicker 
identification than the necessardy long and complicated dichotomic " Key " 
to the species (pp. 90-97). To serve sich purpose is the sole object of (he list 
given above. It has bf-en compiled entirely from material exann'ned bv the 
writer, but a reasonable individual variation in size, rather greater than 
that actually observed, has been presupposed in those species of which tho 
available material has not been considered large enough to show the irua 



iiasy7nallus (Liu-kius), ejmlariiis (New Guinea), formosus (Formosa), 
hi/pomelanus (Indo- and Austro-Malaya), leucopterus (Philippines), 
loochuenshi (Liu-kius), marianims (Mariannes), modiglianii (Engauo), 
vaialis (Christinas I., S. of Java), iiawaiennis (Fijis), ocularis (Ainboina 
group), pselaphon (Bonin Is. ; Volcano Is.), rayneri (Guadalcanar, 
Solomons), .s-«/i/c?(s (Andainans), scapnlatiis (Australia), iongamis (Fijis; 
Tongas ; Sainoas), nalanus (E. Carolines), vanilawemh (Sta. Cruz Is.), 
yapeitsis (VV. Carolines). 

140-149 mm. : — Pf. anratus (Loyalty Is.), cafiiceps (Gilolo group; Sula Is.; 
Sanghir), duhsoni (Celebes), epularius (Ne\T Guinea), hypcnnela'nus 
(Indo- and Austro-Malaya), leu<\q)ftrus {VhiW-^-p'mei), loochoensis (Liu- 
kins), lylci (Siam ; Saigon), modiglianii (Engano), morio (Sumba ; Savu), 
ornaius (New Caledonia), pfelajihon (Bonin Is. ; Volcano Is.), rayneri 
(Guadalcanar, Solomon Is.), samu'eiisis (Siimoas), satyrus (Andnnians), 
scapidaius (Australia), seychellensis (Seychelles), tonganus (Fijis ; Tongas ; 
Samoas), tytleri (Andamans). 

loO-lSQ mm. : — Tt. aritl (Maldives), aferrimus (Bawean I., Java Sea), miraius 
{Loyalty Is.), comorensis (Comoros), conspicjllatus (S.E. New Guinea; 
Australia), gcd.diei (New Hebrides ; New Caledonia), goiddi (Australia), 
lavellanus (Vella Lavella, Solomon Is.), lylei iSAaxn; Saigon), melaiiotits 
i'^cohars), niadicus (Nia.s), niger (Mascarenes). ornafus (New Caledonia), 
pilosiig (Pelew Is.), r/ifus (Madagascar), seychellensis (Seychelles), tonganus 
(Fijis; Tongas; Samoas), /(//"^en (Andamans). 

16(K169 mm. : — Ft. alecto (Celebes; Salayer; Lombok), artel (Maldives), 
aterrimns (Bawean I., Java Sea), chryso-proctus (Amboina group ; Sanghir), 
canspiciUa.Uis (S.E. New Guinea, Australia), giganieus (Ceylon ; India ; 
Himalayas); gmddi (Australia), grandis (Shortland, Bougainville, 
Solomon Is.), livingsionei (Comoros), mclanotus (Nicobars), iiiadicus 
(Nias), ««/r;- (Mascarenes), poliocephalus (Au.stralia ; Trobriand Is.), 
ruhianits (Rubiana, Solomon Is.), riifus (Madagascar). 

170-179mm.: — Pf. ale.rlo (Celebes; Salayer; Lombok), vhrysanchen (Gilolo 
group; N.W. New Guinea), chrysoproclns (Amboina group ; Sanghir), 
cons'picillatus {&.'R.l^evi Gmne&; Australia), (/(^ojji'cMs (Ceylon ; India; 
Himalayas), gouldi (Australia), crandis (Shortland, Bougainville, 
Solomon Is.), inferm^diies (Tenasserim), keyensis (Key Is.), living- 
sionei (Comoros), niger (Mascarenes), rufus (Madagascar). 

180-189 mm. : — J't. cltiryaauchen (Gilolo group; N.W. New Guinea), conspi- 
cillatiis (S.E. New Guinea ; Australia), goiddi (Australia), intermedins 
Temisserim), keyensis (Key Is.), neohihernicus (Bismarck Arch.), paptianns 
(New Guinea), vampyrws edidis (Timor; Savu), vampyrus lanensis 
(Philippines), vampyrus natuinm (Natunas ; Borneo). 

1 9()-199 mm. : — iV. aruensis (Aru Is.), mdanopogon (Amboina group ; Banda 
Is. ; Timor Laut ; Sanghir), neohdiernicus (Bismarck Arch.), papuanus 
(New Guinea), iiffitf;.)^/';;*- /«?!ct(sw (Philippines), vampyrus malaccensis 
(Malay Pen. ; Sumatra), vampyrus nainn<e (N:itunas ; Borneo). 

200-209 mm. : — 7Y. ///fA/wo/jor/o?; (Ainboina group ; Banda Is ; Timor Laut; 
Sanghir), neohihnrnicus (Bismarck Arch.), pjapiianus (New Guinea), 
va/npyrus lanensis (Philippines), vampyrus tnalaccensis (Malay Pen. ; 
Sumatra), vampyrus pluton (Baliv Lombok), rampyrus vampyrus (Java). 

210-220 mm. :—7V. vampyrus pluton (Bali; Lombok), vampyrus vampyrus 

B lO i' 

.(,«f,.' ,.• 

', ' A. The Pteropis hypomelanis group. 

Species. — Sixteen (twenty -six fornas): Pt. hypomelanus (eleven 
subspecies), speviosus, riiimvs, palUims, yr'tsem, sdtyriis, faunulus, 
(idmiralitatum, colonus, soJomonis, brtmnetis, ornaius, anratus, dasy- 
'jtiallus, formosKS, snlni{/er. 

liaiKje. — The Austro- and Indo-Malajan siibrcgions, extending 

rxERorrs hypojjelanfs GEorr. 99 

west to southern Indo-China, north to Formosa and South Liu-kiu 
Islands, east and south-cast to Australia, Central Solomon Islands, 
New Caledonia, aad Loyalty Islands. One species in the Mas- 

General characters. — Skull and dentition unmodified Pteropine ; 
posterior basal ledges of premolars quite short, but distinctly 
marked off from teeth (at least in p\ p^, and pj. Ears moderate 
or short, broad, tip not narrowly pointed; interferaoral short or 
undeveloped in centre ; fur short, adpressed on back, tibia naked 
above, except in certain peripheral species (see below). Colour in 
typical forms not peculiar: dark brownish back, pale mantle, head, 
and underparts, but in all Indo-Malayan, as well as in the 
Formosan and Liu-kiu forms, a pronounced tendency to partial or 
complete suppression of light colours. Sexual differentiation incon- 
spicuous : males without glandular neck-tufts (except Ft. hrimneus), 
but generally with slightly heavier canines and slightly more rigid 
fur of mantle. Size moderate or small (forearm 95-146 mm.). — 
The Mascarene species is in certain respects aberrant. 

Differentiation of species. — Pi. hi/pomelanus, in its full specific 
sense, is the most widely distributed species of the genus ; it covers 
the whole area from New Guinea, through the Moluccas, Celebes, 
and Philippines, to Borneo, Sumatra, Cochin-China, Siam, and the 
Mergui Archipelago, each group of islands having generally its 
own race, but is apparently absent from the southern chain of 
islands from Java, through the Lesser Sunda Islands, to the Banda, 
Key, and Aru Islands, as well as from the Andamans and Nicobars. 
In many of these islands it is however replaced by closely allied 
species, Java being apparently the only large island in the Indian 
Archipelago in which no species of the 7ii/pomelanus group is known 
to occur. The numerous races of Pt. h)/pomelanus differ chiefly in 
the colour of the fur ; those inhabiting the Moluccas (liypomelnmis), 
Celebes {macassaricux), and New Guinea (Zm^cm,?) are unquestionably 
the most ordinary-looking: back dark, mantle and underparts 
pale (buffy), as in a majority of species of Pl4ropus-, passing 
•westward, the most noticeable change in the races is a darkening 
or suppression of all huffy colours, not infrequently combined witli 
a sprinkling of the dark colours with greyish ; to the modifications 
in colour are added in certain islands in the South China Sea 
(N. Natunas: Pt. h. canus; Tarabelan Is.: Pt. h. lepidus) a shght 
increase in the average size of the teeth, and in the extreme south- 
west (Engano : Pt. h. enganus) a well-marked decrease in the 
average size of the individuals. — Pt. speciosns (Snla Is.) and 
Pt. mimiis (Celebes, Philippines) are very similar to Pt. hypomelanvs 
in skull, dentition, and colour of fur, but dift'ering by their com- 
paratively smaller size. 

In the Andamans Pt. Ju/pomelanus is apparently rejdaced by the 
smaller-eyed, longer-furred, and very dark-coloured Pt. sati/rus; 
in the Xicobars by Pl f annul us, which in many respects comes 
rather near to the Engano race (enf/anus) of Pt. hypomelunus, hut 
is smaller, with smaller teeth and longer fur; in Timor by tha 

100 jTKRorrs htpojielanvs ORorp. 

smaller and very pale-coloured Pt. fjrhevs; and in the Eanda 
Islands by the closely allied but darker Pt. jmUidiis. The species 
inhabiting the extreme eastern islands of Austro-Malaya, viz. the 
Admiralty {Pi. admiralltatum) and Solomon Islands {Pt. colonus 
and sohmonis), as well as the single Australian species {IH. 
hruHtu-us), differ from Pt, hypomeJaniis chiefly in smaller size, 
relatively smaller ears, and peculiar colour of the lur, Pt. solomonis 
and brtuuiens also slightly in the distribution of the fur (tibia 
clothed above). 

The eleven species referred to above are in all essential characters 
(both in skull, dentition, and externally) typical representatives of 
the hiipomelanux group. So far as the skull and dentition are con- 
cerned, nearly the same might be said of the four extreme south- 
eastern and northern species, inhabiting respectively New Caledonia, 
the Loyalty Islands, P'ormosa, and the Liu-kiii Islands, but in 
these the external characters are more or less conspicuously 
modified. In Pt. ornatus (New Caledonia) and auratus (Loyalty 
Islands) the general structure of the teeth is unmodified hypo- 
melanine, but the individual teeth are rather heavier, the cranial 
crests and the coronoid process of the mandible therefore stronger, the 
fur is longer and less closely adpressed, the tibiae clothed, the coloiir 
different. In the two northern species, isolated the one in Formosa 
(y^<. /o;-mos((*')and the other in the South Liu-kiu Islands (rt. t/as?/- 
iiiallus), the skull and dentition are perfectly hypomelanine, but 
the fur is long and spreading, extending thickly on the uppcrside of 
the tibiae, the ears small, almost concealed in the fur, the colour 
peculiar. Finally in the single Mascarene representative of the 
grouj), Pt. suhn'ujer (Mauritius, lleunion), not only the external 
characters but also the dentition and skull are modified : size of 
teelh reduced, premolars and molars narrow, skull showing the 
modifications usually found in species with weak dentition (see 
above, p. (33) ; ears small, fur long and spreading, extending 
thickly on upperside of tibia, colour dark brown above and beneath 
with paler collar, size very small. 

Affinities of grotq). — The complete lack of specialization in skull 
and dentition (the Mascarene species excepted), the enormous 
.range of group, its differentiation into many species and local 
races, the fact that it is represented by distinct species in the 
Aiidamans, Nicobars, Formosa, Liu-kius, Solomon Islands, Aus- 
tralia, New Caledonia, Loyalty Islands, and Mascarcnes, — are 
evidence that its origin dates far back in the historj- of the genus. 
Its affinities to the Polynesian 1^1. mariannus type are so extremely 
close, that it is difficult or impossible to draw a hard-and-fast 
line between the two groups. If taken together, they cover 
practically the whole area inliabited by the genus, with exception 
of the Himalayas, India, and Ceylon. 


1 . Pteropus hypomelanus, Temm. 

Pteropns hi/pomclaiius (pt.), Dobson, Cat. Cliir. B. M. p. 57. 

(.Synonyms under the subspecies.) 

DIaijnosis, — Skull typical Ptcnipine. Posterior ledge of p\ |)', 
and p, short, but distinctly marked off from tooth. Ears moderate, 
broad, exposed, tip rounded off. Tibia naked above. Fur of back 
short, adpressod. Pack varying from seal-brown to Proufs brown 
or mars-brown, often conspicuously and sometimes excessively 
sprinkled with silvery greyish-white ; mantle and underparts 
varying from pale buffy, through various darker shades, to chestnut 
or seai-brown. Size below medium: forearm 121-14.5-0 mm. 
Jfah. Indo- and Austro-ifalayan Archipelagos, west to Siam and 
Mergui Archipelago. 

Skull (tig. G, on p. 02). — Typical Pteropine. Deflection of brain- 
caso moderate, alveolar line if projected posteriorly passing through 
middle or upper half of occipital condyles. Itostrura long, rather 
lf)w and slender, somewhat compressed laterally ; distance from tip 
of postorhital process to gnathiou larger than distance from former 
jioint to lambda. Orbits rather large, diameter greater than width 
of rostrum across alveolar borders of p'-p' ; front of orbit vertically 
above some point of front half of m'. Postorbital processes long, 
curved, strong, in aged individuals separated by narrow space from, 
but rarely quite reaching, corresponding processes on zygoma; base 
of postorbital processes raised distinctly above level of frontal 
jtlateau, making this latter between orbits somewhat concave. 
Sagittal crest well developed. Coronoid moderate, somewhat 
sloping; coronoid height of mandible less than c-m.^ ; condyle 
considerably above level of alveolar line of mandible. 

Teeth (fig. 6, on p. Q'l). — Xo special modifications. Ujjper canines 
distinctly recurved ; cingulum moderate, forming a well defined but 
rather narrow rim at inner and posterior l)ase of tooth, p^ minute, 
terete, early deciduous. Posterior liasal ledge of p^ short, separated 
postero-extcrnally by distinct notch from base of outer main 
cusp ; ledge of p' generally less distinctly marked off from tooth, 
m' simple, m" generally slightly larger than, sometimes equal to, 
p,. — i^ twice or twice and a half the bulk of i,. Lower canines 
recurved, cingulum moderate, p, once and a half or twice the size 
of i... Posterior ledge of p^ short, though generally distinctly 
sei)arated from tooth postero-exteriially ; in p, still shorter and 
postero-external notch rather often obsolescent, mj and m., simple, 
in^ equal to or smaller than j),. 

Paliite-nd;/e<i (tig. U, on p. 71).— 5-|-5-|-3, but occasionally with 
some trace of an additional ridge between the normal ninth and tenth 
(compare I'f. auratut). First ridge terminating laterally at front of 
canines ; second at back of canines ; third at front of p' ; fourth 
at back of j)' ; fifth at (middle of) p' ; sixth at front of m' ; 
seventh at back of ra' : eighth at m" ; ninth and tenth behind m" ; 
eleventh to thirteenth situated at palution border. 


Ears. — ;^^ode^ate in length, not nearly reaching eye, brond, 
exposed. Inner margin convex from base to tip, though rather 
ilatter in upper than lower half; outer margin convex in lower 
two-thirds, straight or very flatly concave in upper third. Upper 
half of conch broad, not constricted, tip rather broadly rounded off. 
Conch naked, except posteriorly at base and along lower third or 
half of inner and outer margins. 

Winf/s. — About 15-22 millimetres apart at origin from back. 

Inter femoral. — Very short or undeveloped in centre. 

i'^td-.— Short, adpressed on back. Longest haiis at middle of 
back 10-14 mm. AVidth of hairy space across middle of back 
3S-55 mm. 

Above, humerus covered with short, closely adpressed hairs ; 
elbow nrtked ; thinly spread, short, closely adpressed hnirs on 
proximal fourth or third of forearm. Knee and upper surface of 
tibia naked. Fur extending on lateral interfemoral along inner 
side of tibia to about middle of same. 

Below, antebrachial membrane, lateral membrane along outer 
side of forearm and between humerus and femur covered with 
woolly hair. Tibia naked. 

Colour. — Varying according to the subspecies (see summary of 
modifications of colours, pp. 102-104, and descriptions of subsjiecies). 

liavr/e. — The Indo-Malayan and Austro-Malayan Archipelagos, 
eastward to New Guinea, Trobriand Islands, and Woodlark Island, 
westward to Cochiuchina, Siam, the Mcrgui Archipelago, Sembilan 
Islands, and Engano — No form of Pf. hi/jwwdanus is known from 
Java, the Lesser Sunda Islands, Timor Laut, Key, and Aru Islands ; 
so that the species chiefly covers the Archipelago north of the Java, 
Flores, Sunda, and Banda Seas. In the Andamans (Pt. satj/ms), 
Nicobars (Pt. faimidus), Timor (Pt. f/t-isczis), Banda Islands (Pt. 
pnllidus). Admiralty Islands (Pt. adnnralitatii'ni)., and Solomons 
(Pt. colonns and solomonis), it is replaced by distinct, but closely 
allied, species. 

Subspecies. — The principal characters of the eleven geographical 
races of Pt. lujpomelanns described below may be briefly summed 
up as follows: — 

(1) Size of teeth : — Unmodified in all races, except Pt. h. canus 
and lepidus; in which the teeth average slightly larger than usual. 

(2) General size of animal : — The same in all races, except 
Pt. h. enqamis, which averages smaller than usual. 

(3) Colour of fur : — The general scheme of coloration in Pt. Ju/jk- 
melanus and its principal modifications according to the subspecies 
are these : — 

(a) Back some shade of brown, varying from nearly black, 
through seal-brown, to Prout's brown or mars-brown ; rarely 
these blackish and brownish shades are perfectly uniform, gene- 
rally more or less sprinkled with greyish-white hairs. The 
greater or less amount of pale greyish admixture on back and 
rump is, broadly speaking, a racial character: — In the Papuan 
and Austro-Malrtj-an races {lateus, lu/ponelanus, macassaricvs) the 


pale sprinkling is thin or absent; in the Philippine race (caiiajianus) 
it is usually more conspicuous, though not changing, to any large 
degree, the general dark impression of the colour of the back ; in 
the Mergui Archipelago and Engano forms (i/eminorum, enjanns) 
it is usually so strong as to change the general effect of the colour 
to a grizzled silvery greyish and blackish (brownish); and in the 
races inhabiting the S. China 8ea Islands {cantis, le/ndi's) it is so 
excessive as to make the colour of the back and rump appear 
silverj- greyish-white more or less sprinkled with blackish. The 
difference in the colour of the back between geographically widely 
separated races (luteus contrasted with rjfeminoruni, Ivj pomeJanus 
with cannif, &c.) is verv great, but, all races taken together, there 
is a perfect intergradation from the one to the other. The 
imlividaal variation in the amount of greyish admixture, within 
the same geographical race, is so considerable that in any race 
specimens occur which are indistinguishable from the average of 
neighbouring eastern or western races : in Pt. h. lomesi (Borneo) 
the dark colour of the back is generally moderately sprinkled with 
greyish ; but some specimens are as grey-backed as camts and 
lepidus, while, on the other hand, some individuals of c/avus and 
lepidus are not greyer-backed than pale-col-oured specimens of 
geminomm and engaims. — The general colour of the bade and 
rump is often further modified by a more or less pronounced 
suffusion with mars-brown, russet or buft'y ; in some individuals 
this suffusion is confined to the rutnp and sides of the back, in 
others extending over the whole of the back and rump. It occurs 
sporadically in all races, though apparently more often in the 
western thaTi in the eastern. 

(6) Breast, belly, and fl«,nks :— In the palest -bellied race, 
Pt. h. htieus (New (luinea), the whole of the underparts, except 
throat and flanks, are cream -buff, buff, or ochraceous-buff. Passing 
from New Guinea westward there is a very gradual decrease in the 
pale-coloured area of the underparts combined with a darkening of 
the colour itself. In hif pomdanus (Gilolo grotip) and iiiacassaricus 
(Celebes, iSanghir) the colour of the breast and btelly is deeper in 
tinge, and the dark colour of the flanks shows a tendency to spread 
fco the anal region, hinder belly, and sides of belly and breast. 
In carjaijamis (Philippines), tomesi (Borneo), annectt'its (S. Natunas), 
lepidus (Tambelan Is.), canus (N. Natunas), and condoremis 
(P. Condor, Cochinchina), the bright coloUr has generally darkened 
to golden ochraceous^ cinnamon-rufous, or chestnut, and become 
restricted to the breast and front of the belly, or the breast only, 
or the centre of the breast. Finally, in tjemimrum (Mergui Archi- 
pelago) and eur/C.nus the underparts are wholly dark-coloured, 
sometimes with, but more often without, somD trace of bright 
colour on the centre of the breast. 

(c) The colour of the mantle and head shows the same gradual 
darkening from the eastern to the western races. Generally 
speaking, the mantle and head are similar, or nearly similar, in 
tinge to the bright colour of the breast and belly. In the eastern 

104 PTEROPUS HlTOllEr-ANrs, 

races, the contrast between the pale (bufFy) niautlo and head and 
brownish back is considerable ; passing from New Guinea and the 
Moluccas westward, the colour becomes darker, partly through a 
darkening of the colour itself, partly through a more or less exten- 
sive clouding with chestnut or maroon-chestnut or even chestnut 
seal-brown. While in the eastern races the mantle and head are 
much lighter than the back, this contrast becomes, through 
darkening of the former, much less striking in the central and 
western races ; and in the generally silvery-backed races {canus, 
lepidus) the contrast is reversed, the hazel or chestnut or blackish- 
chestnut mantle being much darker than the light silvery grey 

(4) Phases : — («) Through partial or almost complete suppression 
of all bright colours, as well as of the greyish admixture (if 
normally present in the race), the colour, in certain individuals, 
becomes approximately blackish above and below, though generally 
some trace of the normal brigliter colour (darkened to chestnut 
seal-brown) is observable on the mantle and head, sometimes also on 
the centre of the breast. This " blackish " phase is not rare in all 
western and central races, which in this phase are entirely indis- 
tinguishable from each other; it appears to be absent in the 
extreme eastern, light-coloured races {hi/p<mielanu,s, luteus). — 
(6) A phase characterized by grizzled grey-and-blackish (dark 
" hair-brown ") head, back, rump, Hanks, and sides of breast and 
belly occurs in some western races (observed in (/em inorian, enganus, 
canns, and annectens, but probably occurring in all western and 
central races). It seems to be due in some cases to an extension 
of the greyish sjuinkling (of normal occurrence on back and rump) 
to head. Hanks, and sides of breast and belly, while in other cases 
it appears to be a blackish jihase moditied by heavy sprinkling 
■with greyish. 

Key to the Subspecies of Pteropus hypomelanus. 

a. Crown and sides of head generally similar in 
colour to hack : blackish thickly mixed with 
greyish-white hairs, produciug the general 
effect of ji very dark shade of hair-brown. 
(Individuals occur with back and head more 
or less suffused with russet or wood-brown.) 
* a'. Mantle chestnut more or less clouded with 
■■ seal-brown, rarely as bright as cinnamon- 

rufous. Size normal : forearm 134-137 mm. [p. 106. 

(Mergui Archipelago) Ft. h. c/eminoruni, 

b' . Mantle much lighter ; varying fi'om bright 
hazel to almost ochraceous-buff. Aver- 
aging smaller : forearm 121-133'o mm. [p. 107. 

(Engano) Ft. h. evf/annx, 

h. Crown and sides of head, in the ordinary phase, 
similar to or paler than mantle, varying from 
deep chestnut through several shades of 
ochra,ceous to almost cream-huff. (In some 

PTERoi'us nyroMELANue. 105 

races a blacliish phase occurs, and a phase 
with head, back, flanks, and belly grizzled 
greyish aud blackish.) 

. Breast and belly or at least centre of breast, 
in the ordinary phase, some shade of cin- 
namon-rufous or hazel or rich ocluaceous, 
darkening on flanks aud anal region (often 
also on belly) into dark chestnut cr seal- 
d\ Back usually dark hair-brown. Forearm 

alxjut 13o mm. (Pulo Condor; Cam- [p. 110. 

bodja ; Siani) Pt. h. condorerisis, 

b^. Back usually pvile mouse-grey or silvery 

whitish-grey sprinkled with blackish, 

with or without buHy or brownish or 

ru<set suft'usion. [Compare c-.j 

a'\ Teeth averaging larger ; p^, length 

4'6"-o'l mm. 

a*. Mantle usually hazel or chestnut ; 

back and rump not, or only slightly, 

suffused with buffy. (A blackish 

phase occurs, and a phase with 

head, flanks, and belly grizzled 

greyish and blackish.) Size as lepi- 

(lits. (N. Natunas) Pt. h. camis, p. 113. 

b*. Mantle usually darker, chocolate or 
seal-brown ; back and rump con- 
spicuously suffused with buffy or 
brownish. (Phases as in canits.) 
Forearm 131-139 mm. (Tambelan 

Is. ; P. Aor ; P. Tioman) Pt. h. Ipjndus, p. 115. 

b^. Teeth of normal size ; p-", length 4-2- 
48 mm. Back usually strozigly suf- 
fused with golden ochraceous or 
Front's brown. (Phases as in cantis.) 
Forearm about 130-1335 mm. [p. 116. 

(8. Natunas) Pt. h. annetieiu, 

c'. Back usually seal-brown or approximately 
Front's brown more or less sprinkled 
with light greyish. (Individuals occur 
with the back strongly suffused with 
c-''. Mantle varying from dark maroon- 
chestnut to cinnamon-rufous ; prevail- 
ing colour of back seal-brown. Fore- 
arm 128-142-5 nmi. (N. Borneo and 

coast islands ; Sembilan Is.) Pf. h. totnesi, p. 119. 

</3. Mantle averaging brighter, rich golden 
ochraceous-buil'; back more tinged 
with Prout's brown. Forearm 135- 
1415 mm. (Cagayan Sulu; Philip- [p. 121. 

pines) Pt, h, cfi(/aijamts, 

. Breast and belly, in the ordinary phase, 
averaging paler, tawny ochraceous or pale 
ochraceous or ochraceous-buff ; flanks and 
anal region Sfal- brown or chestnut seai- 
brown (a phase with blackijh underparta 

106 riKRorrs HrroMELAxts gkmixokum. 

occurs, at least ia 7nnr(issan'ri/s) ; back and 
rirnip seal-brown Of Front's brown. [Com- 
pare f'.] 
d-. Underparts tawny ocliraceoiis or pale 

ocbraceoiis ; head and mantJe tftwiiy. 

Forearm ].''>l-14o-o mm. (Celebes ; San- fp. 124. 

gfhir Is. 5 Talaut Is.) I'l. h. mncnssaricua, 

€-. Underparts paler, wold' n buify ; head and 

mantle li^bt oehraceous. Size as lore- [p. 127. 

goinjj. (Gilolo g-roHp) I'i. h. hypomelctnus, 

€ . Palest extreme ; underparts, head, and 
mantle oehraceous - bwff or cream-buff ; 
back and rump from seal-brown to mars- 
brown. Forearm 128"0-l;55>5 mm. (New 
Guinea; Trobriand Is.; Woodlark I.) . . PL h. lnhnm, p. li'8. 

1 a. Pteropus hypomelanus geminorum. Miller. 

? Pteropus nicobaricus (pt., nee Zelelov), J. Audersoii, Cat, Mamm. 

Ind. Mus. pt. i. p. 103 (1881 : Merj^'ui Arch.). 
Pteropus gemiuorum, Miller, Smiths. Misc. Cell. xlv. p. 60 (6 Nov. 

190j : S. Twin I., Mergui Arch.) ; id., Fam. 4" Gen, Bats, p. o8 

? Pteropus celseno (pt., nee Herm.), Maso?i, Rec. Ind, Mus. ii. 

pt. ii. p. 105 (1908 : Mergui Arch.). 

Diagnosis. — Back and head generally dark bair-brown, mantle 
deep chestnut, underparts unitorni blackish (a hairbrown-bellied 
phase occurs). Siee not smaller than usual. Forearm 134-137 mm. 
Hah. Mergui Archipelago. 

Colour. — Two " phases " are represented in the small series 
examined : in most specimens the underparts are almost uniform 
Wackish, in one hair-brown like back and crown. Corresponding 
phases occur iu the allied races of the species. 

{a) Black-bellied phase ( c? ad., rf yg. ad., two $ ad. skins ; 
U.S. N. M. nos. 104461, 62, 63, 65).— Back and rump blackish 
or blackish seal-brown more or less heavily mixed with silvery 
greyish-white or buffy-white hairs, producing the general effect of 
a very dark shade of hair-brown. Individual variation not very 
great, chiefly dependent on the greater or less preponderance of the 
])ale greyish element of the colour ; but even in the darkest 
(blackish hair-brown) individuals the pale sprinkling is quite 
conspicuous. — Breast, belly, and Hanks generally nearly uniform 
blackish or blackish seal-brown inconspicuously sprinkled with 
huffy or whitish-grey hairs ; in one specimen ( 5 ad., 104465, 
with mantle rather paler than usual) the centre of the breast is 
slightly washed with dull russet. — Mantle chestnut more or less 
clouded with (sometimes nearly darkened into) seal-brown ; in one 
specimen (104405) rather brighter, nearly cinnamon-rufous ; colour 
gradually darkening on sides of neck, and passing on foreneck into 
the general blackish seal-brown of the underside. The seal-brown 
clouding of the colour of the mantle is (if present) generally con- 
fined to the tips of the hairs; subapical portion same shade of 


ciniiamon-rufous, base generally blackish. — Crown aii'l sides of 
head similar to back, occasionall}- slighth* suffused with a tinge 
similar to or darker than that of mantle. Throat blackish or 
blackish seal-brown like rest of underparts. 

(b) Hairbrown-bellied phase (5 vg. ad. skin; no. 104466).— 
The peculiar colour of this phase is due to an increase in the 
amount of silvery white hairs on back, breast, and belly. Back 
and rump greyish hair-brown. Breast, belly, and flanks mixed 
greyish white and blackish, the former colour predominating. 
Mantle seal-brown inconspicuously sprinkled with silvery white 
hairs ; sides of neck and foreneck nearly blackish. Head grizzled 
greyish white and blackish as rump. 

Measurements. On pp. Ill, 112. 

Specimens examined. Six skins and skulls, paratypes, from the 
collection of the U.S. National Museum *. 

liamje. Mergui Archipelago (S. Twin Island). 

Type in the U.S. National Museum (no. 104464). 

Pteropus geminorum. Miller; 1903. — Miller separated P/. ^e»n'- 
nor^iin from lepidus on account of its darker colour and smaller 
teeth. But lepidus is an unusually pale-backed and large-toothed 
form. The skull and teeth of r/emino)-inn are in every respect 
indistinguishable from those of most other forms of Ft. h>/po- 
vtehmus, and even from those of lejndus they differ only in average 
characters. The grizzled greyish and blackish colour of the head 
and back, the dull chestnut or chestnut and seal-brown mantle, 
and generally blackish underside give a series of e/eminornm, if 
taken as a whole, a somewhat peculiar aspect, but individuals 
which to the smallest details of colour are similar to an average 
specimen of gcminorum are found in lepidus and caeiayamis. 
Pt. h. c/eminorum is a well-marked geographical race of hyjjo- 
nielanus, but I am unable to draw a "hard-and-fast" line between 
this and allied forms of the species. 

1 b. Pteropus hypomelanus enganus. Miller. 

Pteropus hypomelanus (7iec 7'emm. subsp.), Thomas, Ami. Mus. Civ. 

Genova (2) xiv. p. 106 (April, 1894: Eiigano); ? id., torn, cit, 

p. 6G4 (June, 1895: Si Oban, Mentawei) ; Trouessart, Cat. 

Mavim. i. p. 82 (pt.) (1897: Engano); Willink, Nat. Tijd. 

Kedirl. Ind. Ixv. p. liH (pt.) (1905 : Engano ; ? Mentaweij. 
Pteropus (Spectrum) hypomelanus (pt.), Matschie, Megachir. p, 24 

(1899: Engano; ? Mentawei). 
Pteropus enganus. Miller, Proc. U. S. Nat. Mus. xxx. pp. 822, 824 

(1906: Engano, l*ulo Dua, Pulo Mirbau). 

Diagnosis. — Averaging smaller than any other race of the species. 
Back and head darker or lighter hair-brown with or without russet 
or huffy suffusion ; underside approximately seal-brown, more or 
less sprinkled with grey, and more or less suffused with huffy 

' >-os. 101461-63, 10446.5-07. 

108 rrauorrs HtroMKLAxrs engantts. 

or ochraceous-buft' ; mantle brighter than in (/i^minoruiu, varying 
from bright hazel to almost ochraceous-buti'. Forearm 121- 
13;Vo mm. Hah. Engano. 

Sl-iiU and teeth. — Skull averaging decidedly smaller than in 
fjeniiiiorum and hpieliis, but scarcely distinguishable from that of 
small individuals of cat/ai/ann-'i. Teeth averaging smaller than in 
i/eniiiwnon ; p', length 'd-7—i'6 mm., against 4"2-4'8. Most of the 
teeth are absolutely smaller than in h-piihts. 

External dimoviions. — Pt. h. emjanus averages in every respect 
smaller than any other race of the species ; forearm 1 21-133'5 mm., 
against 128-145-5 in all other races; third metacarpal 82-91, 
against 87-{'7-5 ; lower leg 54-60, against 57'5-(iH ; calcar 1 1-14, 
against 13-18. The sexes do not seem to differ appreciably in 
size; in six adult males the forearm measures 121, 125"5, 12.'i'."), 
]28*5, 129'5, and 133-5 mm., average 127'3; in seven adult 
females 123-5, 126-5, 127, 127-5, 128, 128, and 120, average 

Colour. — rT ad., paratype, teeth slightly worn, 2 Nov. 1904 
(U.S. N. M. no. 140960). Back and rump blackish brown rather 
thickly mixed with silvery greyish white, ])roducing the total effect 
of a dark shade of hair-brown very similar to that of an average 
specimen of r/emi)iormn : rump very slightly washed with a paler 
brown. — Breast and belly blackish brown, thinly but distinctly 
sprinkled with shiny silvery grey hairs. — Mantle bright hazel, in 
some places lightened by golden ochraceous-buff, in others clouded 
with cinnamon-rufous, darkening on sides of neck to chestnut, 
this again on foreneck to dark seal-brown. Base of hairs of mantle 
not differing. — Crown and sides of head light greyish hair-brown ; 
individual hairs silvery whitish grey, everywhere mixed with 
blackish or dark brown hairs ; throat similar in colour to foreneck. 

The specimen described above is one of the most plain-coloured 
in the series, but it scarcely represents the average coloration in 
this form ; in most individuals the back, head, and undcrparts are 
more or less conspicuously suffused with russet or light tawny 
ochraceous. The individual variation may be briefly summed up 
as follows : — 

Back and rump : owing to the greater or less amount of sprink- 
ling with shiny silvery greyish-white hairs, the general colour 
varies from dark hair-brown (pale sprinkling comparatively thin), 
through greyish hair-brown, to mouse-grey. Some individuals are 
scarcely, or only very slightly, washed with russet on back and 
rump ; others are conspicuously suffused with light russet or 
ochraceous-buff along edges of membranes and inner sides of tibije, 
but only slightly so on middle of back and rump ; in others the 
russet suffusion extends over the whole of the back and rump, 
making the central portion mars-brown, the lateral edges along 
membranes tawny ochraceous-buff; one specimen ( J ad., 140968), 
representing the extreme in pale colour, is almost wood-brown 
edged with buffy along membranes. — Breast and belly: in some 
tpecimeus api>roximately seal-brown slightly sprinkled with shiny 


silvery grey hairs ; but the greyish admixture is often so pre- 
dominant as to make the general colour a darker or lighter hair- 
brown clouded with seal-brown in the centre of the breast; in 
other specimens a greater or less amount of the greyish hairs 
have turned buff or ochraceous-biiff, making the total colour a 
seal-brown or hair-brown more or less strongly, and more or less 
extensively, suffused with buff, ochraceous-buft', or light russet. — 
Mantle varying from bright hazel or bright russet through several 
lighter tinges to yellowif-h ochraceous-buff ( $ ad., l-i0968) ; the 
foreneck is always seal-brown or blackish seal-brown. — Crown of 
head : a darker or lighter shade of hair-brown, dependent on the 
smaller or greater amount of grizzling with silvery greyish white ; 
in man)- specimens more or less strongly washed with huffy. 

The variation in colour is independent of the sex of the indi- 
viduals (six males, eight females examined) ; thirteen of the 
Bpceimens were obtained in Xovember, one in July ; teeth in 
different stages of wear (from sliglitly worn to well worn). 

Measurements. On pp. Ill, 112. 

Specimens exiimined. Fourteen (nine skins), from the collections 
of the U.S. Xafional* and British Museums. 

liatiije. Engano, with Pulo ])ua (one mile S.E. of Engano) and 
Pulo Mirban {\\ miles S. of Pulo Dua) ; ? Mentawei Islands 
(no specimens examined). 

Type in the U.S. National Museum (no. 140966). 

Pteropits e»ganiis,^\\\\cY ; lUOl!. — Type localit} : Pulo Dua, off 
Engano. Described by Miller as " similar to Ft. hpidus, but 
smaller, and with back darker than mantle : larger than Ft. hi/po- 
mehtnus, Temm." It is true that in ein/amis the mantle is brighter 
than the back, whereas in the normal, pale-coloured phase of 
lepidus the mantle is much darker than the back ; but lepidua 
is an unusually pale-backed form ; in all races of Ft. hi/pomeJanvs, 
except lepidns and (/eminorum, the back is, as a general rule, 
darker than the mantle, so that in this respect e»(fanus does not 
differ from most other forms of the species. The statement that 
emiauus is larger than luipomelan-us {i.e. Ternate individuals) ie 
based on Dobsou's measurements (forearm 11 'J mm.) of what be 
calls the " type specimen," but which is in reality not the type 
of hifpomelfiims Temm. but of tricolor Gray, and this specimen is 
immature ; as pointed out above, eiu/anus averages, on the contrarv, 
in every respect smaller than any other known race of the species. 

liemark.^. — This form is undoubtedlj- most closely allied to 
Pt. h. (jeminorwn, from the Mergui Archipelago. It accords with 
yemiiiorum in the grizzled light grey and blackish-brown colour 
of the back and head, and the generally dark (seal-brown) colour of 
the l)reast and belh', sprinkled with light grey hairs; but it averages 
smaller, the mantle is noticeably brighter, and there is a much 
stronger tendency in en<iiniui to a russet or buff'y wash of the 
colour of the upper and under side. So far as the available 

» Nos. 1 41 »'.iiW -(•>!', Hii«i()I-7<', l4*)<tT2. HllHta-i, 141007^; paiatvpee. 


material goes, engnmis is alw^ays separable, at least by the brighter 
colour of the mantle, from geminorum, but certain individuals of 
c/emitiorum are iu every respect indistinguishable from some speci- 
mens of lepidiis aud cagai/anus, and the same is the case with 
certain individuals in the series oi enganwa \ in other words, though 
apparently always separable from each other, geminonim and 
enganus are so intimately related to the forms inhabiting the 
islands on the eastern side of the Malay Peninsula as to be some- 
times indistinguishable from these. 

a. J ad. sk. ; skull. Pulo Diia, off En^ano ; Marquis Gr. Doria 
July, ISyi {Dr. E. [P.]. 


1 c. Pteropus hypomelanus condorensis. Pet. 

Pteropus edulis (pt., nee Geoff.), Horsfield, Cat. Mamm. Mus. 

E. Ind. Co. p. 27 (18ol : Siam). 
'• Pteropus condorensis, Peters, MB. Akad. Berlin, 1869, p. 393 

(Piilo Condor) ; Dobson, Mon. As. Chir. p. 18, footnote (1876); 

Matschie, Megachir. pi. viii. tigs. 2, 2rt (skull) (1899 : P. Condor). 
Pteropu'i (Spectrum) hypomdanus g. condoven.sis, Matschie, Mef/a- 

chir. p. 25 (1899 : P. Condor). /. condorensis, Trouessart, Cat. 

Mnmm., Siippl. p. o2 (190i: P. Condor). 
Pteropus nicobaricus (pt., 7iec Zelebor), Dohsoii, Cat. Chir. B. M. 

p. 55 (1878: P. Condor); Trouessart, Rev. Sf Mag. Zool. (3) vi. 

p. 202 (1879: P. Condor); id., Cat. Mamm. i.'p. 81 (1897: 

P. Condor). 
Pteropus hypomelanus {nee Temm. subsp.), Dobson, Rep. Brit. Assoc. 

1880, p." 173 (1880: Cambodja) ; Trouessart, Cat. Mamm. i. 

p. 82 (pt.) (1897 : Cambodja). 

Diagno.n.'i. — Back mixed seal-brown and silvery grey, producing 
the general effect of a dark shade of hair-brown ; breast and belly 
approximately hazel ; mantle blackish chestnut. Size not smaller 
than usual. Forearm about 135 mm. IJab. Pulo Condor ; Cam- 
bodja ; Siam. 

Colour (type). — Back and rump mixed seal-browil and silvery 
grey ; the brownish and greyish hairs about equal in number and 
uniformly mixed, producing the total effect of a dark shade of hair- 
brown ; very faint traces of a pale russet suffusion along edges of 
lateral membranes. — Breast and belh' hazel, brightest (washed with 
golden ochraceous-buff ) in the centre of the breast, gradually 
darkening through chestnut to seal-brown on flanks. Woolly hair 
on underside of membranes blackish seal-brown. — Mantle blackish 
chestnut with hazel bases to the hairs, lightening to chestnut with 
ochraceous-buff hair-bases posteriorly in a transverse line across 
shoulders ; on the sides of the neck and foreneck the colour passes to 
blackish seal-brown. — Upper surface of head chestnut with ochra- 
ceous-buff hair-bases, pas.sing through a darker shade of chestnut 
on sides of bead to chestnut seal-brown on throat. 
"'i Measurements. On pp. Ill, 112. 

Specimens e.vnmined. ]N^ine, in the collections of the Berlin, Paris, 
and British Museums, including the type and six paratypes of the 



BfDu/e. Pulo Condor, off Lower Cochiuchina ; Cambodja ; Siam. 

T>ijje in the T5erlin Museum. 

Fteropiis condorensis, Peters; 1869. — Type: d' ad., teeth well 
worn, mounted, skull separate ; collected in Pulo Condor by 
K. Germain ; lierlin Museum, no. 3945 (from Paris Museum); 
skull figured in ' Megachiroptera des Uerliner Museums,' pi. viii. 
figs. 2, 2 rt. Paratypes : five mounted specimens, same locality and; 
collector, Paris Museum, nos. 46, 46 a (two specimens), 46 bis 
(young), and 46 ter ; and one unmounted skeleton, Paris Museum 
(Mus. d'Anat. Comp.) no. A. 12. 609. Most of the mounted speci- 
mens in the Paris Museum are more or less faded by exposure to 
light ; a skin in the same Museum collected in Pulo Condor by 
Harmand (lleg. no. 30) is similar in colour to the type. 

a. 2 ad. sk.; skull. Si^m (FinlaysoJi). India Musemu [P.]. 

External measurements of Pteropus hypomelanns gemiuorum, 
enganus, and condorensis. 

Ft. hypomelanus 

4 ad. 

14 ad. 


PoUex, total length, c. u 

„ inetiicarpal 

„ 1st phalanx 

2nd digit, metacarpal 

„ 1st phalanx- 

,, 2nd-;5rd phalanx, c. u. 
3rd digit, metacarpal 

,, 1st piialanx 

,, 2nd phalanx 

4th digit, metacarpal 

,, 1st phalanx 

„ 2nd phalanx 

5th digit, metacarpal 

„ 1st phalanx 

„ 2nd phalanx 

Ears, length from orifice 

„ greatest width, flattened . 

Front of eje to tip of mu/.zle 


Lower leg 

Foot, c. u 


66 5 
40 5 





2 ad. 
(Type and 

Mm. Max. Min. Max. Ttpk, Pab.itype.I' 

















16-5 ^ 


































96 5 



























* Approximate measurements (skins). 
f Paris Museum, no. 4tt ter. 



Measurements of sJculls and teeth of Pteropua hypomelanus 
geminorum, enganus, and condorensis. 


Pt. hypomelanus 


Skulls : 4 ad. 

Teeth: 4 ad. 

1 iuiiu. 


: 8 ad. 
9 ad. 



12 2 













5:{ 7 












































25 8 

























17 -8 












„ palatiim to incisive foramina 

„ iVont of orbit to tip of nasals 

„ width of bniin-oase at zjgotuata... 

„ width across canines, externally ... 
„ postorbital constriction 

,, width of mesopterygoid fossa 

,, between p'-p'', internally 

Mandi ble, length 

p', length 

„ width 

„ width 

„ width 

,, width 

I'lKKOl'L'S UYi'OMEI.AXUii CANl'S. 113 

] d. Pteropus hypomelanus canus, A'. And. 

Pteropus hvpomelaiius {vfc Temm. snbsp."), Tliomas, Nov. Zool. ii. 

p. 489 (1896: Pulo Pandak ; P. Pnnjnng; P. Laut) ; Miller, 

Proc. Wash. Acad. Sei. iiu p. 137 (pt.j '(1901 : Pulo Laut). 
Pteropus (Spectrum) hvpoinelaiuia (pt), Mafschie, Megachir. p. 24 

Pteropus hypomelanus cauiis, K. Andeisen, Aim. S)- May. N. U. (8) 

ii. p.36r(l Oct. 1908: N. Natuua.s). 

Diarpiosis. — Teeth averaging larger than in other races of the 
species, except lejndus. Back, in the normal pale-coloured phase, 
jjale, lightening to silvery wliitish-grej" on rump, and 
with or without a huffy suffusion ; mantle some shijdc of hazel or 
chestnut ; head similar to or brighter than raautlc. A blackiah 
phase occurs, and a phase with head. Hanks, and belly grizzled greyish 
and blackish. Size as lepidus. Bab. Xorth Natiiua Is. 

Sl-uU and teeth. — Skull us in Pt. h. cngayanus, but with mandible 
and tooth-rows averaging a little longer. Teeth averaging larger 
than in any other race of Pt. hypomdanKs Gx.ce'pt Upidus: p'*, length 
4'()-5'l mm., against 3'9-4'9 in all other cases exclusive oflepklns; 
p', length 4-7-5'5, against 3' 7-5. 

Colour. — Three colour phases are represented in the small series 
examined, viz. : — 

(1) Back light grey ; head, mantle, and underparts some shade 
of chestnut or hazel or ochraccous ; flanks thinly or heavily mixed 
with greyish (two skins : J ad., type, teeth well worn, P. Pandak, 
Sept., ; $ ad., teeth somewhat worn, I'. Panjang, July, — General colour of back pale mouse-grey, gradualh' 
lightening on hinder back and rump to silvery whitish-grey. 
Individual hairs silvery wiiite and blackish; on tlie anterior half 
of the back the blackish and whitish hairs are nearly equal in 
number, thus producing the general effect of a pale mouse-grey ; 
further back the blackish hairs gradually become scarcer, until on 
the rump the silvery whitish colour is largely predomijiant. All 
hairs with a distinct silky gloss. In one specimen the colour of the 
back and rump is almost imperceptibly, in the other (type) distinctly 
washed with buff'y especially on rump and along edges of membranes. 
— Centre of breast and belly ricli golden ochraceous or orange-buff, 
gradually darkening on sides of breast and bellj- through hazel into 
chestnut ; flanks in one specimen (typ<i) seal-brown thinly sprinkled 
with greyish, in the other pale mouse-grey like back.- — ]Uant]e 
hazel j)artly clouded with chestnut (type), or chestnut -with 
ochraceous-buff'y hair-bases (the other), jjassiiig through a darker 
shade on sides of neck into deep chestnut or chocolate on foreneck.- — 
Crown nearly ochraccous-buff' in centre, darkening through tawny 
ochraccous or deep mars-brown on sides of head into chestnut or 
liglit seal-brown on throat. 

(2) Head, belly, sides of breast, and flanLs grizzled liair-browa 
or broccoli-brown ; mantle and centre of breast sumo shade of 
chestnut ( ? ad., teeth well worn, P. Laut, Sept., 95.1 1 .8.4). — Back 
and rump as in phase (1 ). Belly, sides of breast, aiid flunks a light 
greyish shade of hair-browii rather slightly suff'uscd with dark 


lit PlERul'fS HVl'OMKLAXrS CANU3. 

l)uflV, ])roduch"ig as general effect a shade very nearly approaching 
liroccoli-brown. Mantle dark chestnut, passing through a still 
darker shade on sides of neck into almost seal-brown on foreneck ; 
colour of foreneck continued backward on centre of breast, gradually 
merging into the broccoli-brown of the rest of the underside. 
Crown and sides of head grizzled mouse-grey and seal-brown ; throat 
blackish seal-brown. 

(3) Blackish phase (S jun., P. Laut, Aug., 104737).— Eack 
glossy seal-brown thinly sprinkled with greyish-w^hite hairs particu- 
larly on rump and thighs. The whole of the underside from throat 
to iuterfemoral, including flanks, glossy blackish seal-brown, with 
a faint indication .)f a brighter (glossy seal-brown) tinge on fore- 
neck ; a few greyish-white hairs scattered on breast and belly. 
Mantle dark chocolate. Crown and sides of head dark seal-brown. 
Phase (1), as described above, is chiefly characterized by the 
bright (chestnut, hazel, ocliraceous) colour of the head, breast, and 
belly ; but the flanks may occasionally be thickly grizzled with pale 
greyish. In phase (2) the greyish admixture has spread over the 
head, flanks, belly, and sides of breast, entirely or partly replacing 
the normally brighter colour of these portions of the pelage ; at the 
same time the colour of the mantle and foreneck is darkened to 
chestnut or seal-brown. It is probable that a larger series will 
show a gradual transition from (1) to (2). The blackish phase (3) 
stands apparently more isolated ; it seems to be due to a partial or, 
in some places, total suppression of the greyish element, combined 
with a darkening of the chestnut parts into chocolate (mantle) and 
blackish seal-brown (head and underparts). 
Measuranenis. On pp. 117, 118. 

Specimens exainiiwd. Four, from the collections of the U.S. 
National * and British Museums. 

Range. North Natuna Islands : Pulo Panjang, P. Pandak, P. Laut. 
Type in collection. 

llemarls. — This form belongs to a small group of races of 
Pt. hypomelanus characterized (so far as the normal light-coloured 
phase is concerned) by the strong predominance of silvery white in 
the colour of the rump and back. Three races of this group are 
known. Ft. h. canus, lepidus, and annectens, distributed over probably 
all the islands in the S. China Sea ; in two of these races, canus 
(N. Natunas) and lepidus (Tambelan group, P. Aor, P. Tioman), the 
teeth average rather larger than usual in the species. Pt. h. canus is 
very closely related to lepidus, differing, in its normal phase, only in 
the conspicuously brighter tinge of the mantle and head ; the blackish 
phase is apparently indistinguishable from that of lepidus ; to the 
phase described above under (2) I have seen no equivalent in lejjidits. 

a. 9 ad. sk.; skull. Pulo Panjang, N. Natunas ; Tring Museum. 

July, 1894 {E. Hose). 

6. d' ad. sk. ; skull. Pulo Pandak, N. Natunas ; Tring Museum. 

Si'pt. 1894 (C'h. Hose). {Type <if subspecies.) 

r. 5 acb sk. ; skull. I'ulo Laut, N. Natunas: Tring IVIuseum. 

Sept. 1894 {E. Huse). 

* r^ .ind skull; Pulo, N. Nut.nKi«, Aug. 8, 190a {Dr. W. L. 
AbboH); Reg. no. lOlT'iT. 


1 e. Pteropus hypomelanus lepidus, Mdler. 

Pteropiis lepidus, 3IiUer, Proc. Wash. Arnd. Sci. ii. p. £37, text-fi?. 

(skull) (20 Aug. 1900 : Saddle I. : BigTambelan I.) ; Trouesmrt, 

Cat. Mamm., Suppl. p. 53 (1904 : Tambelan Group) ; Miller, 

Fam. ^- Gen. Bats, p. 68, fig. 8 (skull) (1907). 
Pteropus hypomelanus lepidus, Tliomns, Journ. Feci. Mai. St. Mas. 

ii. no. 3, pp. 101, 102 (1908: Juara Eny, P. Tion;an). 
Pteropus (livpomelanus) lepidus, Kluss, Journ. Fed. Mai. St. 3Ius. 

ii. no. 3, p. 153 (1908: P. Tioman ; Permangil ; Aor). 
? Pteropus nicobaricus (nee Zelebor), Bonhote, P. Z. S. 1900, p. 875 

(Apr. 1901: Great Iledang I.); Kloss, Journ. Fed. Mai. St. 

Mus. ii. no. 3, p. 152 (1908 : (treat liedang I.). 

Diagnosis. — Similar to Ft. li. canns, but mantle and head, in tho 
normal pale-coloured phase, conspicuously darker. Forearm 1!^1- 
139 mm. Hah. Tambelan group ; P. Aor; P. Tioman. 

Skull and teeth. — As in Ft. li. canns. 

Colour. — (1) Pale phase (six skins). — Light greyish colour of 
back sometimes nearly without any trace of butfy suffusion (one 
specimen, J ad., Sept., P. Tioman;; generally strongly 
washed with golden buffy (one, (j" ad., Aug., Big Tambelan I., 
101649), or irregularly clouded with ])ale cinnaraon-buffy or very 
pale mars-brown (three; $ ad., Aug., Big Tambelan I., 101650; 
5 ad., June, P. Aor, 112404; d ad., June, P. Tioman,— 
Centre of breast golden ochraceous tinged with orange-buff, or rich 
huffy russet, or chestnut-hazel, passing into dark chestnut on sides 
of breast and centre of belly ; flanks greyish broccoli-brown or light 
hair-brown sprinkled with blackish hairs, this grizzled colour some- 
times extending to sides of belly and anal region. — Mantle sometimes 
chocolate with paler hair-bases, biit generally more or less clouded 
with seal-brown, sometimes nearly uniform seal-brown ; sides of neck 
and head, foreneck, and throat similar or darker. Crown similar 
to mantle or paler. 

(2) Blackish phase ( J ad., Aug., Big Tambelan I., 101651).— 
Indistinguishable from corresponding phase of Ft. h. caiias. Tlie 
only specimen has a faint trace of a brighter colour on the centre of 
the breast. 

Measurements. On pp. 117, 118. 

Specimens examined. Seven, from the collections of the U.S. 
National* (three. Big Tambelan T., paratypes ; one, P. Aor), Kuala 
Lumpur (one, P. Tioman), and British Museums. 

Eancje. Tambelan Islands (Saddle I. ; Big Tambelan I.) ; Pulo 
Aor ; Pulo Tioman ; probably also Permangil and Great Kedang 
Island (see references above). 

Type m the U.S. National Museum (no. 101670, Saddle I.). 

Itemarlcs. — The closest known relative of this form is undoubtedly 
Ft. h. canus ; but in the colour of the back (light greyish generally 
more or less strongly suffused with huffy or some buffj' shade of 
pale brownish) it shows leanings towards its eastern neighbour, 
Ft. h. anneciens, from S. Natunas. 

* X.J3. JU1(149-:.I; 1121114. 

IIG PTEKorrs HYPoaiErANVs annecteks. 

«. d nd. sic. ; Jiiara Bay, Pulo Tioiaan, S. China Kuala Lumpur S.l.i?5.2. 

skull. Sea; '14 June, liX)6 {H. C. Museum [P.]. 

h. J !\(1. sk.; Juara Bay, 'P. Tioman ; 14 Sept. Kuala Lumpur 

skull. 1907. Museum [P.]. 

1/. Pteropus hypomelanus annectens, K. And. 

rteropus hypomelaiuis {nee Tcmnt. siibsp.), Tliomas Sr Hartevt, Nor. 

Zool. i. p.' 655 (1894 : Sirhassen) ; Miller, Proc. Wash. Acad. Sci. 

iii. p. 137 (pt.) (1901 : Sirhassen). 
Pteropus (Spectrum) livpomelanus (pt.), Matschie, Mcjachiv. p. 24 

Pteropus bvpomelanus annectens, K. Andersen, Ann. !<c Mag. N. H. 

(8) ii. p.'^Gl (1 Oct. 1908 : Sirhassen). 

Dicu/nosis. — Teeth not averaging larger than usual. Normal 
pale-coloiired phase rather similar in colour to corresponding phase of 
Ft. h. lepidas, but generally more strongly suffused with golden ochra- 
ceous or paler or dai'ker Prout's brown on back, and with brighter 
mantle and breast. Forearm \'60-\'6',yo mm. Hah. S. Natuna Is. 

Slcull and teeth. — Only two skulls of perfectly adult specimcn.s 
have been examined ; they average smaller than in Pt. h. canus and 
lepidus (total length 61-;3-64-5 mm., against 62-5-(;)7-2 in canufi 
and Jepid^(s), one of them being in fact quite as small as the smallest 
skull of Pt. h. emjanus. Teeth (five specimens) as in most races of 
the species, not averaging larger than usual as in canus and lepidus : 
p^ length 4-2-4-S mm. (in Pt. h. tomesi 4-2-4-8), against 4-0-5-1 
in canus and lepidus. 

Colour. — Three colour phases are represented in the series 
examined, viz. : — 

(1) Pale phase with ochraceous, or cinnamon-rufous, or mars- 
brown underside (three specimens ; imm. al., 94 9.28.25 ; 
cJ ad. skin, teeth slightly worn, June, 104732 ; cJ jun. skin, 
June, 104733). — Back and rump a light shade of hair-brown 
approaching mouse-grey and very strongly suffused with golden 
ochraceous or paler or darker Prout's brown, particularly along 
middle of back and rump; greyish fur distinctly sprinkled with 
blackish hairs. — Breast and belly ochraceous darkening to cinna- 
mon on flanks (, or uniform cinnamon-rufous slightly 
clouded with chestnut and with concealed ochraceous-buff bases 
to the hairs (104732), or approximately dull mars-brown, palest 
and washed with golden buffy in centre of breast, darkening to 
seal-brown on flanks (104733). — ^Mantle generally closely similar 
in tinge to coloi;r of underparts : ochraceous, gradually darkening on 
sides of neck, passing into hazel on foreneck (, or hazel 
clouded with chestnut, darkening to chestnut on foreneck (104732), 
or dark mars-brown, passing to dark vandyck-brown on foreneck 
(104733). — Crown similar to mantle or paler, darkening on sides 
of head into colour of throat, which is similar to that of foreneck. 

■ (2) Grey -bellied phase (one ; $ ad. al., teeth well worn, — Similar in colour to corresponding phase of IH. h; 
canus (supra, p. 113, pluise 2). 

(3) Seal-brown-bellied phnso (one ; S jun, skin, 104734).— Back 



and rump cssontially as in the ordinary pale ])liase. Centre of breast 
seal-brown, darkening to blackish seal-brnwn on sides of breast, 
belly, and llauks. Mantle dark chocolate, passing to seal-brown 
on sides of neck, foreneck, throat, and sides of head. Crown slightly 
paler than mantle. — This specimen probably represents some transi- 
tional stage rather than the true " blackisb phase "' ; the whole of 
the nnderparts is, rongbly speaking, dark seal-brown, but the colour 
of the upperside does not show any clear sign of darkening. 

Measurements. I5elow and on p. 118. 

Specimens examined. Pive, from the collections of tie U.S. 
National * and British Museums. 

Bancfe. South Xatuna Islands : Sirhassen. 

Type in collection. 

KijmarJiS. — In characters as in habitat this race seems to occupy 
an intermediate position between Pt. h. lepidus and iomesi ; in the 
size of the teeth it agrees with the latter form, in the colour of the 
fur it approaches the former ; in the size of the skull it may prove 
to average smaller than either. 

a. Iiiiui. al. ; skull. 

b. 2 ad. al. ; skull. 

Sirhassen, S. Natunas. 

A. Evnrett [C.]. 

( Ti/pc of subspecies.) 
A. Everett [C.]. 94.9.2S.20. 

External measnrements q/Ttcropus hypomelanus canus, lepidus, 
and anncctcns. 

Koi-ea rtu 

PoUes, total length, o. u 

„ metacarpal 

,, 1st phalanx 

■Jnd digit, metacarpal 

,, 1st phalanx 

2iid-3rcl plialans, c. ii. 

3rd digit, metacarpal 

,, Ist phalanx 

,. 2ik1 phalanx 

-1th digit, metacarpal 

,, 1st phalanx 

,, 2nd phalanx 

5tL digit, metacarpal 

„ 1st phalanx 

,, 2nd phalanx 

Ears, length from orifice 

,, greatest width, flattened .. 

Front of eye to tip of inu/zle 


Lower leg 

Foot, c. u 


Pt. h//poinelaims 

3 ad. 
(Incl. type.) 








94 o 












14 5 














7 ad. 

2 ad. 

Mi.v. Max. Mis. Max. 



































































* A'os. 104732-34; Sirhassen. 

t Approxiai.'ito measurements (skins). 



Measurements of sJculIs and teeth of Pterojms hypomelanus canus, 
lepidus, and annectens. 

Ft. hypo 



Skulls : 3 ad. 
Teeth : 3 acl., 

1 imm. 
(Incl. type.) 


Skulls and 

teeth : 

6 ad. 


Skulls: 2 ad. 

Teeth: 2 ad., : 

3 imm. i 

(Inch type.) i 

Skull, toUl length to gn nth ion 









































30 2 























24 2 


























Max. j 

mm. 1 

64 5 1 

22 S 


7'7 1 









3 1 

2-8 i 


22 ' 


.5 1 

3 1 

5 t 



29 1 

3 8 




„ palation to incisive foraiiina 

„ front of orbit to tip of nasals 

„ width of brain-ease at zygomata ... 
,, z.vgoitiatic width 

„ width across m', externally 

„ lachrymal width .... 

,, width across canines, externally... 

,, width of mesopterygoid fossa 

,, between p'^-p', internally 

„ between cingula of cauiries 

,, orbital diameter 

Mandible, leno-th 

„ coronoid height 

Upper teetli, c-m^ 

Lower teeth, c-m, 

Upper incisors, combined width 

p', length 

„ width 

p^, length 

,, width 

ni', length 

,, width 

m^ length 

,, width 

„ width 

D.. lensth 

„ width 

„ width 

■t^Awr. ■ IL :»^,iivi !)t<r- 

riEUorrs iiypojiklaxi's xoaiksi. 119 

!(/. Pteropus hypomelanus toinesi, Fet. 

? Pteropus vocifei'us, Peale, U.S. E.vpl. E.vp. viii. Mamm. p. 19, 

text-ti;;. (skull) (1848: Mangsi 1., Balabac Straits); Warner, 

Schrebers Sduij., ib'uppl. v. p. 001 (1853-uo: Mangsi) ; Giehel, 

Sdug. p. 1003, footnote (IHua : Maiio-si) ; Fitzimjer, SB. Akad. 

Wien, Ix. Abth. i. p. 444 (1870: Maugsi). 
? Pteropus (Spectrum) hypomelaiiusy. vociferus, Matschic, M((/achir. 

p. 25, c/'. p. 11 (1899: ^langsi). e. vociferus, Trouessart, Cat. 

Mamm.,Siipi)l. p. 52 (1904: Maiigsi). 
? Pteropus inacliloti [nee Temm.), Cassin, U.S. Hxpl. Exp. viii. 

(2 ed.) p. 10, text-lig. (skull) (1858: Maagsi); Feters, MB. 

Akad. Berlin, 1867, p. 333 (pt.) ; Dobson, Cat. Chir. B. M. p. 00 

(pt.) (1878) ; Trouessart, Bev. S,- Ma//. Zool. (3j vi. p. 203 (pt.) 

(1879) ; id., Cat. Mamm. i. p. 83 (pt.)"(1897). 
Pteropus hvpomelanus (iicc Temm. subsp.), Tomes, P. Z. S. 1858, 

p. 536 (1859: Labuau); Dobson. Cat. Chir. B. M. p. 58 (pt.) 

(1878 : IJorneo) ; Trouessart, liev. ^- Maq. Zool. (3j vi. p. 202 

(pt.) (1879: Borneo); Hose, Mamm. Borneo, p. 38 (1893); 

Everett, P. Z. S. 1893, p. 494 (Borneo) ; Trouessart, Cat. Mamm. 

i. p. 82 (pt.) (1897: Borneo); Willink, Nat. Tijd. Nederl. Ind. 

Ixv. p. 274 (pt.) (1905 : Borneo). 
Pteropus hypomelanus var. toniesi, Peters, MB. Akad. Berlin, 1868, 

p. 620 (Sarawak). 
Pteropus (Spectrum) hypomelanus c. tomesi, Matschie, Megachir. 

p. 25 (1899: Borneo), d. tomesi, Trouessart, Cat. Mamm., Suppl. 

p. 52 (1904: Sarawak). 
Pteropus (hypomelanus) tomesi, Kloss, Journ. Fed. Mai. St. Mas. 

ii. no. 3, p. 153 (1908 : Pulo Kampia, Sembilau Is.). 

Diaijnosis. — Similar in size of teeth and external dimensions to 
Pt. h. annectens, but on the whole much darker ; ajiprosimately seal- 
hrown thinly sprinkled with greyish, and with mantle, head, and 
breast varying from dark raaroou-chestnut to ciiiuamon-rufous. 
Forearm i23-112-5 mm. Ilab, N. Borneo, with adjacent coast 
islands ; Serabilan Is. 

Skull and teeth. — Teeth as in Pt. h. annectens, but the skull may 
prove to average a little larger. 

Colour.- — Ordinary phase : breast a pale shade of hazel or 
cinnamon-rufous, generally washed with orange-bufF (four speci- 
mens ; ad. skin, teeth slightly worn, April, Mengaluu I., 
o ad. al., teeth slightly worn, Darvel Bay, ; d ad. skin, 
teeth much worn, July, Sibutu I.,; ad. skin, teeth some- 
what worn, Aug. or Se])t., Lamboyan I., — Back and 
rump varying from blackish seal-brown to pale seal-brown, slightly 
or moderately, but never heavily, sprinkled with silvery whitish 
hairs. — Breast, or at centre of breast, a pale shade of hazel or 
cinnamon-rufous, as a rule more or less conspicuously washed with 
orange-buff; on sides of breast, flanks, and belly the colour 
gradually, sometimes rather abruptly, darkens to seal-brown or 
blackish seal-brown with or Avithout a slight admixture of greyiah 
hairs. — Mantle varying from dark maroon-chestnut to che.stnut 
cinnamon-rutous or chcstnul-hazcl, becoming gradually darker on 


sides of neck, and dark chestnut or seal-brown on forencck. — Crown 
similar to mantle, or rather brighter, or somewhat clonded with 
dark brown, indistinctly sprinkled with greyish or buffy hairs, 
passing througli a darker shade on sides of head, into blackish seal- 
brown on throat. 

A fifth specimen ( cJ ad, al., teeth slightly worn, Darvel l)ay, 
1) is similar to the above in the colour of the mantle, head, 
and underparts, but on the sides of the back the number of silvery 
greyish-whito hairs, washed with buffy, has so greatly increased as 
to change the general effect of the colour to butfy greyish-white 
sprinkled with blackish ; rump greyish-white suffused with buffy 
and russet and slightly sprinkled with blackish. In the colour of the 
back and rump this specimen comes very near to certain individuals 
of Pt. h. lepidus, the only noteworthy difference being that the 
grevish-whitc colour does not extend to the median portion of 
the back. 

In a sixth specimen (J ad. al., teeth much worn, Darvel Bay, 
94,7.14.1) the whole of the back is very strongly suffused with 
russet, darkest on front of back, gradually merging into a paler 
(more golden ochraceous) shade on rump, and the colour of tlie 
breast and belly is rather brighter, more washed with ochraceous, 
ihan usual. In tho whole of the coloration this specimen comes 
very near to Pt. Ti. annectens. 

The blackish and the grey-bellied phases of this form are as yet 

Measurements. On pp. 125, 126. 

Sjieclmens examined. Eight, iu the collection of the British 

E'(i)ge. Sarawak ; Labuan and Mcngalun Islands (off N.W. 
Borneo) ; Sibutu Island (cff N.E. Borneo) ; Lamboyan (off S. 
TalaAvan) ; Scmbilan Islands (off N.E. Sumatra). 
?'v^)« not in existence (?). 

P^a-op^s )'oc(yVr(;,s, Pealc ; 184S. — T}-pe locality, Mangsi Island, 
Balabac Straits ; type iu the U.S. National Museum (no. 3961). 
Ten years later identified by Cassin (l.s.c.) with Pi. [i. e. Acerodon^ 
macMoti. — The following details about the type-specimen were 
kindly contributed by l)r. Marcus W. Lyon, Washington : Once 
mounted and on exhibition, then put away in storage ; no skull 
found in tho skin, and none in the collection known to belong to it 
[but must have been examined by Peale, since figured by him] : 
specimen not full-grown, actual length of forearm 117 mm., of 
tibia 60 mm. ; furred area of back of normal breadth ; fur so 
bleached that there is no telling what were the original colours, 
— Without an examination of the dentition it is impossible to decide 
whether Pt. vociferus is a Pteropus^ (in which case there can pre- 
sumably only be question of a form of the hijpomehtnus type) or a 
small Acerodon. When to this it is added that the specimen is 
young and the colours unreeognizalile, Pt. vnnfem.s must, for the 
present at least, be put down as indeterminable. 

Pteropus Injpomelanus var. tomesi, Peters: 1868. — Name proposed 

PTKUorrs iivro.MEi.Axcs tojikst. 


by Peters (Z. s. c.) in a nominal list of Cliiroi»tera obtained by tbe 
Manjuis G. Doria in Sarawak. Peters refers, without description, 
to Tomes's paper in P. Z. IS. for 1858 (I. s. c), in which this latter 
author describes a specimen of Ft. hiipomelunus obtained by James 
Motley in the island of Labuan and, in 1858, in the possession of 
Mr. L. L, Dillwyn. Type locality therefore Labuan, and type the 
specimen described by Tomes and considered by him a " dark 
variety " of hypomdanus. 1 have been unable to trace the fate nf 
this specimen ; judging from the descrijjtion given by Tomes in 
was perfectly similar to the specimens here referred to Ft. h. 

lianarks. — Ft. h. tomesi is closely related to Ft. h. ccinus, Jepulua, 
and annectens; it agrees with these races in the colour of tho 
mantle, head, and underparts ; but whereas in canus, Icpithcs, and 
annectens the back and rump are silvery greyish or silvery whitish 
sprinkled with blackish and with or without a buffy, russet, or 
Prout's brown suffusion, these parts are in tomesi, owing to a 
decrease of the silvery whitish and corresponding increase of the 
dark clement of the colour, blackish or seal- brown sprinkled with 
greyish. The intimate relationship between all these races is 
further proved by the fact that tomesi is occasionally almost indistin- 
guishable ill the colour of the fur from lepidus (tbe fifth specimen 
described above) or annectcns (the sixth). In dentition tomesi 
agrees perfectly with the majority of races of hypomelanus, i. e. it 
does not show the enlargement of the average size of the teeth 
characteristic of crtnus and lepidxis. 

a-c. 2 c? ad., 12 ad. al. 

d. Ad. sk.; skLiU. 

e. c5"ad. sk. ; skull. 

/. cT;id. ,nl. ; skull. 
g. Ad. sk. ; skull. 

h. Oad. fk. ; skull. 

D.arvel Bay, N.E. Commander of 

Borneo. H.M.8. 'Egeria' 

Mengalun I., off A. Everett [C.]. 

N. \V. Eorneo ; 

April, 1.S92. 
SibutuL.cffN.E. A. Everett [C.]. 

Borneo ; July, 

Sibutu I.; Julv, A. Everett [C.l. 

Lambovan I., off DougLos Cator, 

S. 'Palawan ; Esq. [C. & P.]. 

Aug. or Sept. 

Pulo Rampia, Kunla I'.umpur 

Seunbilan Is., iluso-jni [P.]. 

N.E. Sumatra ; 

7Aug. 190G(i/. 

C. Rob in SOU)), 

1 h. Pteropus hypomelanus cagayanus, Mearm. 

Pteropus [.sp.], Walvrhoiisc, P. Z. 6'. 1S4.3, p. G" (Philippines). 
Pteropus hypomelanus (wee Tciivn. subsp.j, Dob.wn, Cat. Chir. B. JT". 

p. 58, .specimens «-fi? ( l.'^rB : Din.igat) ; Troncsxarf, licv. S: Maq. 

Zool. (3) vi. p. 202 (,pt.) (1870 f S.E. Philippines); Glinther, 


p. Z. S. 1879, p. 74 (Dinag-at ; Sariyao) ; Steere, List Birds 4- 

Mamm. Steere Exp. Pkilipp. p. 28 (1890: Guimaras ; Paiiay ; 

Leyte) ; Elera, Cat. Sist. Faun. Filipinas, p. o (pt.) (I8U0 : 

Luzon ; Dinagat ; Mindanao) ; Elliot, Field Col. Mus. Puhl. ii. 

Zvol. Ser. i. no. 3, p. 76 (1896 : Panay ; Guimaras) ; Trouessart, 

Cat. Marinn. i. p. 82 (1897 : Dina<?at) ; Sanchez, An. Soc. Espan. 

Hist. Nat. xxix. pp. 241,275,288(1900-1901: Luzon; Leyte; 

Panay ; Guimaras) ; Elliot, Cat. Mamin. Field Col. Mus. p. 491 

(1907: Panay). 
Pteropus (Spectrum) liypomelanus (pt.), Matschie, Meijachir. 

pp. 23, 24 (1899: Ciiyos Is.; Mindanao); Trouessart, Cat. 

Mamm., Suppl. p. 52 (1904 : Philippines). 
Pteropus cagayanus, Mearns, Pmc. U.S. Nat. Bins, xxviii. p. 433 

(13 May, 1905: Cagayan Sulu) ; Miller, Fam. i^- Gen. Bats, 

p. 58 (1907). 

Biagnosk. — Similar iu skull, teeth, and external dimensions to 
Ft. h. iomesi, but mantle averaging considerably brighter, and back 
and rump usually of a more Front's brown tinge. A dark-coloured 
phase occurs. Forearm 135-141-5 mm. Hab. Cagayau Sulu ; 
Philippines genei'ally. 

Colour. — (1) Brighter phase: breast and belly, or at least 
centre of breast, golden ochraceous or ochraceous-butf, with or 
without a tinge of orange-buff; mantle similar (three specimens ; 
ad. skin, teeth much Avorn, June, Cagayan Sulu,, topo- 
type ; cJ ad. al., teeth very sliglitly worn, Dinagat, 77. 10. G. 8 ; 
5 yg. ad. skin, Jan., Panay, 105445). — Back and rump varying 
from a dark shade 01 Prout's brown approaching seal-brown to 
almost typical Prout'.s brown, sometimes with a rather slight wash 
of dark russet; practically uniform, or more or less conspicuously 
sprinkled with silvery greyish-white hairs. — Breast, or breast and 
belly, varying from rich golden ochraceous to a tinge intermediate 
between i3utf and ochraceous-buff, in any case generally washed 
with orange-buft'; on sides of breast and belly passing into dark 
chestnut, this again darkening into the seal-brown of the flanks. — 
Mantle rich golden ochraceous-butF, generally tinged with orange- 
bufF and more or less clouded with tawny or hazel, j)articularly in 
front near occiput and posteriorly near back ; passing through a 
darker shade on sides of neck into the dark cinnamon-rufous or 
pale chestnut of foreneck. — Crown of head similar to mantle, or 
more clouded with brownish and grizzled with silvery hairs ; 
passing through a gradually darker tinge on sides of head, into 
seal-brown or chestnut seal-brown on throat. 

(2) Intermediate stages.— One skin ( cJ ad., teeth very slightly 
worn, Jan., Panay, 105441) has the whole of the back nearly 
uniform Prout's brown, and the mantle nearly uniform chestnut, 
lightening to rich golden oohraceous-bufF in a transverse band 
between shoulders. — In two skins ( $ iram., Jan., Panay, 105446- 
47) the bright colour of the underparts is restricted to a narrow 
space on the centre of the breast; mantle approximately as in 105441. 

(3) Dark phase: underparts seal-brown; a narrow space in 
centre of breast washed with tawny or ochraceous; mantle very 


dark ($ imm. al., Diiiogat, 77.10. G.9; d ad. skin, tteUi some- 
•what worn, Jau., Pauay, 105442). — Back and rump as in ordinary 
]ihase. Underparts nniform seal-brown, darkening to blackish on 
throat, and with a small area in centre of breast sprinkled with 
deep tawny hairs (one specimen) or washed with ochraceous (the 
other). Mantle chestnut heavily clouded with seal-brown, gradu- 
ally darkening into the uniform seal-brown of the underparts. 
Head nearly similar to mantle, or strongly grizzled with hair- 

Measurements. On pp. 125, 12G. 

Specimens examined. Thirteen, from the collections of the U.S. 
National t and British Museums. 

ItaiKje. *Cagayan Sulu; Philippines: *Luzon, Cuyos Is., *Panay, 
Guimaras, Leyte, *I>inagar, *Mindanao (Surigao). Specimens 
have been examined from the localities marked with an asterisk. 

Type in the U .S. National Museum (no. 125289). 

Fteropus cagayamis, Mearns ; 1905. — Original description based 
on three skins and skulls from Cagayan Sulu. Separated by 
Mearns from Ft. hypomelanus chiefly on colour characters ; the 
fikins "were temporarily preserved in formalin, but the colors 
[Mearns writes] have not been materially changed." A comparison 
of Mearns's two paratypes with a topotype in the British Museum 
proves, beyond doubt, that the colour of the mantle in Mearns's 
specimens has been considerably bleached by the preserving 
fluid, which has also partly discoloured the membranes ; these 
specimens have therefore been entirely excluded from the above 
description of tho colours. According to Mearns, " the skull and 
teeth of P. cayayanus are decidedly smaller than in Steere's 
specimens from Panay, which he called ' Pteropus hypomelanus ' " ; 
I am unable to see any difference in the size of the skull and 
teeth between three specimens from Cagayan Sulu and five of 
St;eere's specimens from Panay. 

Remarks. — Pt. h. cagayanus is intermediate between tomesi 
(Borneo) and macassarlcus (Celebes) ; from either of these races it 
can only be discriminated on average colour characters. It di&'ers 
from tomesi in the rather brigliter mantle and rather more Front's 
brown tinge of the back ; in both of these respects it agrees with, 
or comes very near to, macassariciis, in which, however, the under- 
parts average slightly paler. 

t'. h. ^ ad., (5 pull. al. Liuou (//. Cicmliiy). Zool. Soc. Col). Kot reg. 

c,d. jad.,; Dinagat. Purchased (Hig- 77.10.68,9. 
skull.^. gins). 

e. Opull. al. Surigao. Purchased (Hig- 


/. Ad. si;.; skull. Cagavan Sulu ; June, Douglas Cator, 05.7.3O.2. 
181)4. Esq.[C.&P.]. 

t Nos. 12o290-91 (Cagavan Sulu : paratvpcs of Pt. mynmnus Mearns) • 
105141-42, 105415-47 ;Panay ; Steere Exi..).' 

124 rxEEorrs htpomklants rjACAssAKicrs. 

1 i. Pteropus hypomelanus luacassaricus, HemJe. 

rteropus hypomelanus {nee Temm. subsp.), Dobson, Cat. C'hir. B. M. 

p. 58, specimens /, g, h (1878: Sanghir Is.; N. Celebes); 

Troiiessart, Hev. <^- May. Zool. (3) vi. p. 20:2 (pt.) (l879 : Sangliir; 

N. Celebes) ; Jentink, Notes Leyden Mks. v. p. 173 (1883 : N. 

Celebes) ; id., Cat. Ost. Mamm. p. 262, specimens /, o (1887 : 

Siao, Sanpfliir) ; id., Cat. Syst. Mamm. pp. 148, 149, specimens 

»(, q, r (1888 : Siao, Sanghir) ; id., in U'ehcr's Zool. Erycbn. 

lieise Niederl. Ost-Ind. i. p. 96 (1890-91 : Macassar) ; Troue'ssart, 

Cat. Mamm. i. p. 82 (pt.) (1897: N. Celebes); Willink, Nat. 

Tijd. Nederl. Ind. Ixv. p. 274 (pt.) (1905 : Saugbir; Celebes). 
Pteropns (Spectrum) hypomelanus (pt.), Mutschie, Meyachir. 

pp. 23, 24 (1899 : Sanghir ; Gorontalo ; Macassar) ; Troiiessart, 

Cat. Mamm., Suppl. p. 52 (1904 : Sanphir; Celebes). 
Pteropus macassaricus, Hcude, Mem. Hist. Nat. Emp. Chin. iii. 

p. 177, footnote, pi. v. lig-. 4 (teeth) (1896 : Macassar). 
Pteropus (Spectrum) hypomelanus i. macassaricus, Matschie, Meya- 

c/iir. p. 26 (1899). h. macassaricus, Troiiessart, Cat. Mamm., 

Suppl. p. 52 (1904 : Macassar). 

Diagnosis. — Similar to Ft. Ji. cagayanvs, but underparts pnler. 
A dark phase occurs. Forearm 131-1 45-5 mm. Hah. Celebes; 
hjanghir Is. ; Talaut Is. 

Colour. — (1) Bright phase : underparts some shade of ochra- 
ceous (four speciraeus ; S imm., S ad. skins, teeth slightly worn, 
Sanghir Is.,, li ; ^ ad. ah, teeth somewhat worn, N. 
Celebes, ; S juv. skin, Talaut Is., — Back and 
rump between seal-brown and Prout's brown, with or without a 
distiiicb wash of russet, and generally rather thinly sprinkled 
with silvery greyish-white hairs. — Breast and belly yellowish ochra- 
ceous-butt', in one specimen clouded with tawny, passing into seal- 
brown or chestnut seal-brown on flanks and anal region ; extreme 
bases of bright-coloured hairs dark brown. — ^lantle tawny, nearly- 
pure in tinge or somewhat clouded with hazel, generally distinctly 
paler posteriorly near back, becoming slightly d^irker on sides of 
neck, and passing into russet, or russet washed with hazel, on 
foreneck. — Crown similar to mantle or rather paler (one specimen), 
or very distinctly grizzled with silvery grey, darkening on sides of 
head into the daik vandyck-brown or seal-brown on tliroat. 

(2) Dark phase: breast, belly, and flanks uniform blackish seal- 
brown ( c? ad. skin, teeth well worn, Oct., Meuado, 
— Back and rump uniform glossy seal-brown. Breast, belly, and 
tianks uniform glossj' blackish seal-brown, irantle rich golden 
tawny hazel tinged with orange-ochraceous, darker on sides of 
neck, and clouded with seal-brown on foreneck. Head similar to 
mantle, but considerablj- clouded with dark chestnut, this latter 
tinge predominating on sides of head, and passing into chestnut 
seal-brown on throat. — The fur of this specimen is new and 
perfectly unabraded. 

Measurements. On pp. 125, 126. 

Specimens examined. Xine, in the collections of the Leyden and 
British Museums. 

liamji', Celebes (Meuado ; Macassar) ; Sanghir islands ; Talaut 
Islands (Lirong). y 



Tiipe presumably in ihe Zi-ka-wei Museum, near Shanghai. 

rtero2>us maccissaricus, Heude ; 1896. — From the description 
clearly a member of the Pt. hypomelanus group. One of ten 
specimens examined by Heude had " le dos d'un ronx fonce ainsi 
que la poitrine et le ventre," i. e. belonged to the dark phase. 

Remarks. — In the skull, teeth, size, and colour of the head, 
mantle, and back, this race is similar to the Philippine form, Pt. h. 
cuffaijamis, but the undcrparts average rather paler. In this latter 
point it shows decidedly leanings to its eastern neighbour, Pt. h. 

N. Celebes. 

a. cJaJ. al. ; skull. 


o ad. st.; skull. 
5 ad. sk. in al. 

Menado, N. Celebes ; 

Oct. 189.5. 
Macassar, S. Celebes ; 

-2'd Aug. 1895. 
Sanghir Is. 

, e. c? imin., (^ ad. 
sks. ; skulls. 
/.; skull. Lirong, Talaut Is 
May, 1897 {J. 

Dr. A. B. Mever 

Dr. Chas. Hose 

Drs. Sarasin[E.]. 

Dr. A. B. Meyer 

Hon. W. Roth- 
schild [P.]. 

72.4. 15.a 

External mensurements of Pteropus hypomelanus tomcsi, 
cagajanus, and macassaricus. 

Fortsiri n 

Pollex, total lenglli, c. u 

,, metacarpal 

., 1st plialanx 

2ud digit, metacarpal 

„ Istphalans | 15 

,, 2nd-3rd phalanx, c. u. .1 14'.5 

3rd digit, metacarpal j S75 

,, 1st phalanx j ('i3 

,, 2nd phaLinx ' 94 

•Itb digit, mctiicarpal ' 85-.5 

,, 1st phalanx o3.5 

2nd phalanx : 54-5 

otli digit, metacarpal ' 91 

,, 1st piialanx 39 

., 2nd phalanx ' 41 

Kars, length from orilice 245 

„ grcate-st widtb, H.ittened ... 17 

Front of eye to tip of niuz«le ' 24 


Lower leg ' 60-5 

Foot, e. u 4<^r5 

Calcar 14-5 



Measurements of skulls and teeth 0/ Pteropus hj-pomelauus tomcsi, 
cagayanus, and macassaricus. 

Ft. hypometanus. 

Skulls and 
teeth: 7 ad. 


cagayanus. | macassaricn. 

Skulls: 5 ad. 

Teeth: 6 ad., 

4 imm. 

MiK. Max. ! Min. Max. 

Skull, total length to gnatbion 

,, palation to incisive foramina ... 
,, front of orbit to tip of nasals ... 
„ width of brain-case at zygomata 

„ zygomatic width 

t „ width across ni', externaliy 

„ lachrymal width 

,, width across canines, externally. 

' ,, postorbital constriction ' 

,, interorbital constriction i 

j „ -width of mesoptei'vgoid fossa ...\ 

! „ between p*-p*, internally [ 

■ „ between cingula of canines \ 

j „ orbital diameter I 

Mandible, length 

„ coronoid height | 

I Upper teeth, c-m'"' | 

1 Lower teeth, c-m3 1 

j Upper incisors, combined width .. 

I p'', length 

„ width 

p*, length 

I ,, width 

I m', length 

,, width 

! in^, length 

I „ width 

p, , length 

,, width 

i p,, length 

I ,, width 

I p.,, length 

I ,, width 

j in J, length 

j ., width 

i m.^, length 

I ,, width 

1 m^, length 

I ,, width 













































































































Skulls: 3 ad. 

Teeth: 3 ad., 

2 imm. 

MiN. Max. 



































7 6 



























1 j. Ptevopus liypomelanus hypomelamis, Temm. 

Pteropua hypomeliinus, Tenuninck, Esq. Zool. p. 61 fpt.) (1853 : 
Ternate) ; Wagyier, Schreher's Siiug., Suppl. v. p. 599 (1853-55; 
Ternate) ; Peters, MB. Akad. Berlin, 18(57, p. 330 (Ternate) ; 
Fitzinger, SB. Akad. Wien, Ix. Abtla. i. p. 434 (1870 : Ternate) ; 
Dobson, Cat. Chir. B. M. p. 58 (pt., viz. speciiueii i) (1878 : 
Ternate) ; Trouessart, Kei: Sr Mag. Zool. (3) vi. p. 202 (pt.) 
(1879 : Ternate) ; Jejitink, Cat. Ost. Mamm. p. 2()1 (pt., viz. 
specimens b-e) (1887: Ternate); id., Cat. 6'gst. Mamm. p. 148 
(pt., viz. specimens c, d, f, g, h) (1888 : Ternate) ; Trouessart, 
Cat. Mamm. i. p. &2 (pt.)"(l.'>97 : Tcnmte) ; Willinh, Nat. Tijd. 
Nedert. Ind. Ixv. p. T>\ (pt.) (1905: Ternate; Gilolo) ; Miller, 
Fam. ^- Gen. Bats, p. 58 (pt.) (1907). 

Pteropua (Spectrum) hypomelanus, Matschie, Megachir. p. 23 
(pt.) (1899: Ternate); id., Abh. Sencke?ib. Ges. xxv. Ileft ii. 
p. 208, pi. xiii. figs. 4, 4 a (skull of younir) (1900 : Gilolo ; milk- 
dentition) ; Trouessart, Cat. Mamm., Suppl. p. 52 (pt.) (1904 : 

Pteropus tricolor, Gray, Cat. Monk. ^c. p. 108 (1870 ; Ternate). 

" Pteropus spectrum, Matschie," Willink, Nat. Tijd. Nederl. Ind. 
Ixv. p. 274, among synonyms of Pt. hypomelanus (1905) (probably 
error for Ptei-optes {Sixctrum) h/pomelamis). 

Didf/nosis. — Intermediate in colour between Pt. h. jnacassariaix 
and the race of the species, Fl. h. luteus (New Guinea). 
Size as these races. Ilah. Gilolo group. 

Colour. — 2 imm. skin, Ternate, 60.^.26.1, tj-pe of Pt. tricolor. 
Gray.— Back and rump pale seal-brown with a few shinv greyish 
white hairri. — and belly glossy golden bufi'y ; extreme base 
of hairs dark brown ; flanks and anal region dark vaudj'ck- brown. 
— ilantle ochraecoiis-bufF tinged with oehraceou.s, passing into a 
darkei- shade of ochraceous on sides of neck and forencck ; extreme 
base of hairs dark brown. — Crown whitish grey slightly sprinkled 
with blackish hairs (old, somewhat abraded fur), and irregularly 
blotched with a colour similar to that of the mantle (new fur, on 
the point of replacing the mixed whitish-grey and blackish hairs); 
bides of head grizzled greyish and brownish ; throat dark brown. 

A dealer's skin (Verreaux), labelled Ternate, closely resembles 
ihe above in colour, but the crown is light ochraceous without 
admixture of whitish grey. 

Measurements. On p. lol. (P'xtornal measurements omitted, 
owing to immaturity of specimens examined.) 

Specimens e.vamined. Seven, in the collections of the Le^'den 
(five) and British Jluseums, including the cotypes of the species 
and the typi' of Pt. tricolor, Gray. 

Bam/e. Ternate; Gilolo. 

Cotypes in the Leyden Museum. 

PteropxtH hypomelanus, Temminck ; 185;^. — Based on " jilusieurs 
individus '■ collected in Ternate, by Forsten. Of these are now in 


the Leydeu Collection three mounted specimens (Cab. Syst. p. 148, 
l>, c, d) and two skulls (Cat. Ost. p. 261, a and e). An exami- 
nation of these specimens has satisfied me that tho original 
material on which Temminciv founded Ft. liypomelanus consisted 
of two different species, Ft. hupomelanus and caniceps. The 
mounted specimen h {S ad.) and tho skull a are caniceps ; the 
former is, no doubt, the specimen to which Temminck himself 
called attention as differing in colour from the others (Esquisses, 
p. Q2). The real cotypes of lujpomelanus are, therefore, the 
mounted specimens c and d, and the skull e (not full-grown). 
Cotype c is a 2 imm., not quite full-grown ; cotype d \s a, S yg- ad., 
quite or nearly full-grown, though with unconsolidated iinger- 
joiuts. The colour of the fur is in both somewhat bleached, but 
has undoubtedly been quite similar to that of the British Museum 

Fteropus tricolor, Gray ; 1870. — Type locality, Ternate ; typo 
(skin and skull) in collection. Based on a topotype of Ft. liypo- 
vielanus. The species called by Gray " Ft. hypomeJas " (Cat. 
Monk. &c. p. 110 ; 1870) is Ft. araensis, Pet. 

«. 2 imm. sk. ; skull. Ternate. Dr. A. B. Wallace [C.]. fiO.S.-2r..l. 

( 2v//)e of I'f. iricolor, Gray.) 
b. Imm. st. ; skull. Teniate. Tomes Coll. (Verreaux). Not, reg. 

' 1 1-. Pteropus liypomelanus luteus, A'. And. 

Pteropus hvpouielanns (nee Temm. suhsp.), Dobsnn, Cat. Chi?\ 
JJ. M. p. 58, specimen j (1878 : S. of Plunn Gulf, New Guniea) ; 
Trouessart,EeD.^- May. Zool.{^)\'i.^. 20-2 (pt.) (1879: HiioiiGulf) ; 
Dobnon, P. Z. 8. 1878, p. 875 (1879 : Amberbaki, New Guinea); 
Peters i^- Doria, Ann. Mus. Civ. Genova, x\i. p. ()90 (1881: New 
Guinea); Phomas, Nov. Zool. ii. p. 163 (180o : Fergiisson I., 
D'Entrecasteaux Ciroup) ; id., oj). cit. iii. p. 526 (1896: Kiriwiiia 
I., Trobriaiul Group) ; id., Ann. Mus. Civ. Genova, (2) xviii. 
p. 608 (1897 : AVoodlark I.) ; Heller, Ahh. Mus. Dre.^den, vi. 
no. 8, p. 4 (1897: AmberbaUi ; Iluon Guif ; Trobriand : D'Entre- 
casteaux) ; Prouessart, Cat. Mamvi. i. p. 82 (pt.) (1897: New 
Guinea) ; Willink, Nat. Pijd. Kederl. Ind. Ixv. p. 274 (pt.) (1905: 
New Guinea). 

Pteropus (Spectrum) hvpomelanus, Matiichie, Megachir. p. 24 (pt.) 
(1899: Mafur, N.W. New Guinea; Woodlark I.) ; Prouessurt, Cat. 
Mamm., Siippl. p. 52 (pt.l (1904: N.W. New Guinea). 

Spectrum hypomelanum, MatscJiie, in Kriexjer^ Neu-Gninea, p. 77 
(1899: N. New Guinea); Jentink, Notes Leydcn Mvs. xxviii. 
pp. 1G4, ^09 (1906). 

Pteropus hvpomelanus luteus, K. A)ulersen, Ann. if Mar/. N. II. (8) 
ii. p. 362* (1 Oct. 1908: New Guinea; Contlictls. ; Trobriand Is. ; 
AVoodlark I.). 

Diitf/ywsis. — The palest race of the species: back, rump, and 
flanks some shade of brown (from nearly seal-brown to mars-brown); 
head, mantle, throat, forcneck, breast, and belly some shade of 
ochraeeous-buff, buif, or cream-buff, with or without a brownish 

n-KJtoM? jjvi (i.\ji;i.ANys lltkis. 129 

^Aaeh on throat aud anal region. forearm 12<S-,j-13o'r) mm. 
Hah. New Guinea, as far south-east as Woodiark I. 

Colour (whole series examined).^ — Back and rump varying in 
tinge from a sliade of brown approaching seal-brown (two speci- 
mens), through vandyck-brown (two), to mars-brown (three), the 
lighter tinges, viz. vandyck-brown and mars-brown, evidently due to 
a more or less complete suffusion of a darker shade of brown with 
russet (as is the case in most races of the species), a slighter suffusion 
with russet making the general effect of (he colour vandyck-brown, 
a stronger suffusion mars-brown. Colour uniform, or rather paler 
on rump thnn on back. The usual admixture of silvery greyish-white 
hairs in all specimens so slight as not to change the general uniform 
aspect of the colour. — Breast and belly varying froin ochraceous-butf 
with a peculiar golden hue, to cream-buff, this colour in most 
specimens extending over the whole of the breast and belly, while 
in others the liinder part of the belly is more or less (in one 
strongly) clouded with brown of a shade similar to that of the back. 
Bright-coloured hairs uniform from tip to base, or with concealed 
dark brown bases. Flanks brownish, similar to back. — €rown, 
mantle, sides of neck, and foreneck similar to breast and belly ; 
sides of head often washed with russet or brownish ; throat similar 
to foreneck, or clouded with paler or darker vandyck-brown. 

No blackish phase known. 

Measurement!^. On pp. 130, 131. 

Specimens examined. See list below. 

liange. New Guinea; Conflict Is. (Itamarina) ; Trobrjand Group 
(Kiriwina) ; Woodiark I. 

'J't/jie in collection. 

liemarks. — The difference in colour between Pt. h. luteits and any 
of the western races ((/eniiiiu^i, em/nnus, condorensis, cantis, lepidw., 
anneciens, tomest) is great, but the gap is completely over-bridged 
by those races which, step for step, through the Gilolo group {hj/po- 
melanus), Ctilehes (macassaricus). and Philippines (rar/f'y/rtMMs), lead, 
in colour as in geographical habitat, from hiieas to the darker- 
coloured western races. The intimate relationship between all 
these forms is further shown by the fact that, save iu the colour 
of the fur, they are in all respects (». €. in skull, teeth, ears, 
(juality, distribution and length of fur, and external dimensions) 
indistinguisliable from each other, except enganvs which averages 
smaller, aiul ran us and lepidu^ in which the teeth average slightly 


c? yg.ail., c? 
iiiiiii. kI. ; 

2*ew Guinea. 

i?<^v. S. Maofarkiie 




Ad. sk.; 

S. of Huon Gulf, 
New Guinea. 

Dr. P. Comrie [C.]. 


S iiuii). sk. 

Itamarina, Con- 
flict Is. ; 17 Jan. 

W. Stalker [C.j. 

Not reg. 


c. Iram. sk. ; 


/. c? id- sk. 

g. 2 ad. al. 



A. S. Meek [C.]. 

Kiriwina I., Tro- 

briand Group ; 

15 Feb. 1895. 
Kiriwinal.; 17 Feb. 

Woodlark I. ; Mar, 

1890 (Dr. L. 



A. S. Meek [C.]. 

{Type of subspecies.) 
Museo Oivico, 

Genoa [P.]. 

External measurements of subspecies of 
Pteropus hypomelauus. 

Pi. hi/pomelanus 


4 ad. 

(Incl. type.) 


PoUex, total length, c. u 

„ metacarpal 

„ 1st phalanx 

2nd digit, metacarpal 

„ 1st phalanx 

,, 2nd-3rd phalanx, c. u. 
3rd digit, metacarpal 

„ 1st phalanx 

„ 2nd phalanx 

4th digit, metacarpal 

„ 1st phalanx 

,, 2nd phalanx 

5th digit, metacarpal 

,, 1st phalanx 

,, 2nd phalanx 

Ears, length from orifice 

„ greatest width, flattened . 
Front of eye to tip of muzzle .... 


Lower leg 

Foot, c. u 


MlN. M.4X. 

51 -o 





All races, except 

enganus *. 

40 ad. 

MiN. Max. 























* For comparison with measurements of Pi. 
smaller average size of that race. 

enganus (p. Ill), to show 



Measuremeiita of slculls ami teeth of xubspecies of 
Pteropus h3-pome]anus. 

Pt. hypomelanus 

hypomelamis. \ lu/eiis. 

Skull: I ad. I Skulls: 4 ad. 
Teeth: 1 ad., Teeth: Sad., 
2 iinm. 2 imm. 

All races, except 

canus and 

lepidus t. 

Skulls : 3.5 ad. 

Teeth : 3(J ad., 

14 i>iim. 

SbiH, tot'il lengtli to gnathion 
,, palation to iiipisive foramina. 
., irout of orbit to tip of nasals . 
width of brain-caseat zygomata 

zygomatic width 

width across m', externally ... 

lachrymal width 

width across canines externally 
,, postorbital constriction 
interorbital constriction 
width of mesopterygoid fossa 
between p'-p^, internally 
,, between cingnla of canines . 
orbital diameter 

Mandible, length 

„ coronoid height 

I'pper teeth, c-m'^ 
Lower teeth, c-m^ 
L'pper incisors, combined width 

p ', length 

,. width 

p^, length 

., width 

ui', length 
„ width 
in^, length 
„ width 
p,, length 
„ width 
p,, length 
,, width 
p,, length 
,. width 
ni,. length 
,, width 
ni... lengtli 
„ width 
: m,, length 
! ., width 

* Senile specimen. 

t-For comparison with measurements of P^ k. caniHinnd lepidus (p. 118), to 
show, slightly larger avpi-acrp size of mandible, upper and lower tooth-rows, and 
some of the clieekteetli in those races. 






3-5 i 


3 1 


2r, 1 




2 1 






















13ti PTi;Rorus speciosus. 

2. Pteropus speeioBus, K. Ami. 

Pteropus speciosus, K. Andersen, Ann. (^ May. N. II. (8) ii. p. 364 
(1 Oct. 1908 : Malanipa I. ; Sibutu I.). 

Biar/nosis. — Similar to Ft. hypomelanus, but with considerabl_v 
Bmaller skull. Back blackish sprinkled with light gre)- ; breast ami 
belly approximately orange tawnj-. A blackish phase occurs. 
Forearm about 120-123 mm. Hab. Sulu Archipelago. 

ShuU and teeth. — Skull similar to that of Ft. hi/pomelanns, but 
considerably smaller: total length about 57 mm., against 61-69 in 
all forms of hypomelanus ; median palatal length (palatiou to 
incisive foramina) 28'5-29, against 30-35 ; mandible 45'2-46"2, 
against 48-55. — Individual teeth verj- nearly of the same size 
as in small-toothed specimens of the smallest race of hi/pomelanns, 
Pi. h. enffanus. 

Colour. — Light i)hase : (^ ad. al., teeth slightly worn, Malanii)a 
I.,, type. — Hack and rump blackish, conspicuously 
sprinkled all over with shining silvery whitish-grey hairs, producing 
the general effect of a grizzled blackish and pale grej'. — Breast and 
front of belly orange-tawny, somewhat clouded with chestnut 
laterally towards the flanks ; extreme base of hairs dark brown. 
Flanks, hinder belly, and anal region similar to back. — Mantle rich 
hazel, becoming rather lighter and more tinged with ochraceous 
posteriori)' near back, and passing through a darker shade on sides 
of neck into chestnut on foreneck.- — Crown buffy slightly sprinkled 
with blackish hairs ; sides of head seal-brown sprinkled with buflfy 
grey ; throat blackish thinly mixed with buffy grey (total aspect 
similar to that of flanks). 

Dark phase : 2 ^d. sk., teeth extremely worn, July, Sibutu I., — Back and rump seal-brown somewhat varied with 
light grey, particularly on sides of back along membranes. Under- 
parts seal-brown, washed with dull russet on breast. Mantle 
nearly vandyck-brown, becoming darker on sides of neck, gradually 
passing into the daik seal-brown of the foreneck. Crown and 
sides of head grizzled light grey and blackish seal-brown, producing 
the total impression of a dark hair-brown ; throat blackish. 

The lighter phase described above is rather similar to that of 
Ft. h. tomesi, though with the colour of the mantle more like that 
of Ft. h. cagayaniis ; the dark phase is almost indistinguishable in 
colour from the black-bellied phase of Ft. h. r/eminorum. 

External dimensions. — As the smallest specimens of Ft. h. 

Measurements. On pp. 134, 135. 

Specimens e.ramined. Two, in the collection of t)ie British 

Rnngf.. Sibutu I., offN.E. Borneo; Malanipa I., off Zamlioangy. 

Type in collection. 

liemarls. — The skulls of l.he Sibutu and Malaoipa speftirtit>ns, 
one from each island, are perfectly alike; the teeth in the Sibutu 

acl. al. ; 

jralaiiipal., off Zain- 


boaiiga (' Chal- 

lenger ' Exp.]. 

ad. sk. ; 

Sibutu I., off N.E. 


Borneo ; Julj', 



skull are so excessively worn as to be useless for study ; the Sibutu 
plan is evidently in the blackish, the Malanipa specimen in the 
light phase. 

Lords of the Treasury 

{Type of species."* 
b. $ ad. sk.; Sibutu I., off N.E. A. Everett [C.]. 

3. Pteropus mimus, K. And. 
Fteropus Jujpomelanus (pt.), Dobson, Cat. Chir. B. M. p. 57. 

Pteropus hypomelanufj (nee Temm.), Dobson, op. .?. c. p. 58, specimen 
e (1878 : Luzon) ; A. B. 31eyer, Abh. Mus. D?-esden, vii. no. 7, 
p. 6 (pt.) (1899: M.acassar). 

Pteropus mimus, K. Andersen, Ann. ^- Mag. N. II. (8)ii. p. 364 
(lUct.l9U8: Macassar j. 

Diagnosis. — Similar to Pt. speciosns, but back uniform vandvck- 
browii and underparts much lighter. A blackish phase occurs. 
Forearm about 127 mm. Nah. Celebes ; Philippines. 

Colour. — 2 ^^- skin, teeth much worn, Macassar,, type: 
Back and rump vandyck- brown washed with mars-brown oa rump 
and with seal-brown on interfemoral along inner side of femur and 
proximal half of tibia. — Jireast aud belly pale golden ochraceous 
tinged with orange, heavily clouded with mars-brown on breast and 
crissum, purer in tinge on belly ; concealed base of fur dark brown. 
Flanks dark Prout's brown, many of the hairs with tawny tips. — 
Mantle rich cchraceous-buff strongly tinged with orange, palest 
posteriorly near back, darkest (more tawny) in front ; sides of neck 
and foreneck nearly tawny. Concealed bases of hairs seal-brown. 
— Crown similar to mantle; forehead and sides of head brownish 
mixed with buffy hairs or tips to the hairs; throat seal-brown. 

A second skin from Macassar ( c? juv., not full-grown, 
is similar to the foregoing, but mantle considerably darker, between 
cinnamon and russet. This specimen is of particular interest as 
showing an initial transitional stage to the dark phase, the faded 
and abraded pale brownisli fur on the rump being replaced by fresh 
blackish seal-brown hairs. 

A si)ecimen from Luzon (' J ad. al., teeth well worn, colour 
apparently not deteriorated, is similar to the type, but 
with breast lighter, more deep golden ochraceous-butf. 

The colour of these specimens is much like that of Pi. ht/pouichtnus 

Measurements. On pp. 134, 13-5. 

Specimens examined. Three, in the collection of the British 

Ran(je. Celebes (Macassnr) ; riiilippiucs (Luzon). 

Tijp': ia collection. 



Ihmarls. — This species is similar to Fl. sj^eciosus in skull and 
iceth, but differs considerably in the colour of the fur. I am 
unable to distinguish the single Luzon from the single adult 
Macassar specimen. — Of six specimens collected by the Drs. Sarasin 
in Macassar and mentioned by Dr. A. B. Meyer (l. s. c), one is 
evidently young (forearm 100 mm.); if the others are adult they 
probably belong to this species (forearm 120-128 mm.). 

a Pad. al.; skull. Luzon. Dr. A. B. Meyer 

/*. 2 ad. sk.; skull. Macnssar (i)r. .-i. /?. Tomes Coll. 

Wallace). ( Type of species. ) 

c. c?; skull. Macassar ^i);-. .1. /i'. Touies Coll. 


External measurtmenU of Pteropus speciosus and 

Pi. speciosus. > I't. mimus. 

2 ad. I 2 ad. 

(Type and paratype.) ! (Type and paratype.) 

6 ad. 





Pillex, total length, o. u 

,, 12 

1st phalanx | -'8 _ 

2nd digit, metacarpal ! 59 o 

I ., J st phalanx | 1^5 

! ,. 2nd-3rd phalanx, c.ii., ^^ 

; 3rd digit, metacarpal , °' _ 

1 ,, 1st phalanx ' *''"^ 

' ., 2nd phalanx \ 89 

i 4th digit, metacarpal 

I ,, 1st phalanx 

I ,, 2nd phalanx 

1 nth digit, metacarpal 

,, 1st phalanx 

2nd phalanx 

Eai-s, length from orifice 

„ greatest width, flattened 
Front, ol' eye to tip of muzzle . 


Lower leg 

Foot, c. u 


2 ad. 



















35 5 











2 ad. 
















2 ad. 
Paratype t, 


127 "5 

















* B. M. 

+ B. M. 



Measnrements of skiiUs and teeth of Ptcropus speciosus 
and mimiLS. 

Type a 

nd paratype. 

Ft. mimus. 

Skulls : 2 ad. 

Teeth : 2 ad., 1 imm. 

(Inch type.) 

Skull, total length to gnatliion 

„ palation to incisive foramina. 

„ i'ront of oi-bit to tips of nasals 

,, widtb of brain-case atzvgomata 

zygomatic width ....! 

6 ad. 








































































„ width across mi, exterually ... 
„ lachrymal width 


„ width across canines externally 

„ postorbital constriction 

„ interorbital constriction 

„ widtb of mesopterygoid fossa. 

„ between p'-p*, int'enially 

„ between cingula of canines ... 
orbital diameter 




Mandible, length 


,, corouoid height 


Upper teeth, c-m- 


Lower teeth, c-m, ... 


Upper incisors, combined width 

p^, length 



„ width 


p', length 


„ width 

m^ length 

' ,. width 


1 m', length 


„ width 


p,, lengtli 


j ,, width 


P3, length 


„ width 


P41 length 


„ width 


m,, length 


' „ widtli 


'. ni., length 


1 ,] width 


nij, length 


„ width 


iy6 - fifiRoprs pALLiiitts. 

4. Pteropiis pallidus, Tonm. 
Pteropus griseiis (pi.), Dobson, Cat. Chir. B. M. p. 44. 

Pteropus pallidus, Temminck, Mon. Mannn. i. p. 184, pi. x^-. figs. 8, 
9 (sluill) ('1825 : Banda Is.) ; Lesson, 3Ian. Mamm. p. Ill, no. 288 
Q827 : Banda) ; Lesjivtrest, Diet. Sri. Nat. xlvi. p. 363 (1827 : 
Banda) ; Is. Geofroi/^ Diet. Class. d'Hist. Nat. xiv. p. 701 (1828 : 
J3anda) ; J. B. Fischer, Syn. Mamm. p. 84, no. 9 (1829 : Banda) ; 
Lesson, Ilixt. Nat. Mamm. (Comj^l. Bvffon) v. p. 54 (1836: 
Banda) ; Temminck. Mon. Mamm. ii. p. 77 (1b37 : Banda ; 
Sumatra and Malacca, errore) ; Gray, Griffith'' s Anim. Kingd. 
V. p. •'iOS (1838 : Banda) ; Oken. Ally. Natury. vii. Abth. ii. p. 990 
(1838); Wagner, Schrebefs Sciuy., Suppl. i. p. 352 (1839: 
Baudrt) ; S. MiiJler, in Temminck' s Nat. Gesch. Nederl. Overz. Bes., 
Zooyd. pp. 20, 59 (1839-44 : Banda) ; Lesson, N. Tabl. R. An., 
Mamm. p. 13, no. 182 (1842 : Banca, errore) ; Schinz, Syst. Vers. 
iS'rtur/. i. p. 126 (1844: Bauda) ; Gray, Xoo!. ' Samarang,' Vert. 
p. 12 (1849 : Banda) ; Wagner, Schfehefi Stmy., Siippl. v. p. 600 
^•,, V (1853-55: Banda): Gervais, Hist. Nat. Jfrtwrn. i. p. 187 (1854 : 
t.'l Banca, erroie); Giehel, Siiug. p. S98 (1855: Banda); Finsch, 
Neu-Guinen, p. 150 (1805 : Banda); Fitzinger, SB. Akad. Wien, 
Ix. Abth. i. p.' 43G (1870: Banda). 
Pteropus (Spectrum) hypomelanus c. pallidus, Matschin, Megachir. 
p. 24 (1899 : Brtuda). b. pallidus, Trouessart, Cat. Mamm., 
-■ ■ Suppl. p. 52 (1904 : Banda). 

Pteropus grireus (nee Genf.), Peters, MB. Akad. Berlin, 1867, 
p. 326 (pt.) (Banda) ; Dobson, Cat. Chir. B..M. p. 44 (pt.) (1878 : 
Banda) ; Trouessart, Rev. Sr May. Zool. (3) vi. p. 205 (pt.) (1879: 
Banda); Jentink, Cat. Ost. Mamm. p. 255 (1887: Banda); id. 
Cat. Syst: Mamm. p. 141 (1838 : Banda) ; Trouessart, Cat. Mamm. 
i p. 80 (pt.) (1897 : Banda); Mafschie, Meyachir. yA. i. iigs. o, 
ba,bb (skull) (1899: Banda). 
Prevopus hypomelanus (pt., nee Temm.), Jentink, Cat. Ost. Mamm. 
p. 261, specimens/-/ (1887: Banda); id. Cat. Syst. Mamm. 
p. 149, specimens w-z (1888 : Banda) ; Trouessart, Cat. Mamm. i. 
p. 82 (1897: Banda). 

Jhaqnosis. — Similar in .skull and teeth to Pt. mimnx, but breast 
aftd belly dark brown (seal-brown) strongly grizzled with pale hairs, 
external dimensions smaller. roi-earm 113*5-1]9 mm. Hah. 
iSanda Is. 

Colonr. — Approximate description from two Leyden cotypes, 
fnonnted, somewhat faded (Jentink, Cat. Syst. p. 141, Pt. griseus^ 
f,. ]iY — Back, rump, breast, belly, and flanks seal-brown or dark 
brown strongly sprinkled with greyish, oi* whitish-grey, or buffy- 
grev hairs, thus producing the "couleur de feuille morte'' mentioned 
bv Temminck. Mantle, sides of neck, and foreneck ochraceous-butt' 
(faded?). Head buffy (faded?) ; chin and throat dark brown like 
breast and belly (much faded in " a "). Hairs of head, mantle. 
sides of neck, and foreneck with concealed dark brown bases. 

The third LoA-den cotvpe {'' r" ) i*^ more uniform dark brown 

MiciSutements. Ou pp. 140, 141. 

Specimens examined. Nine, viz. five cotypes of species in the 
Lej'clen and Berlin Museums, and four mounted speeimens with 
BkuUs in the Leyden Museum, collected in Pulo Ai, Banda group, 
by Hoodt (Jcntink, Cat. Syst. p. 149, sub Pt. l^jpomelanus, w, x, 
y, z; Cat. Ost. p. 261, Pt. hypomelanus, f, g, h, i). 

Panr/e. Bauda Islands. 

Cotypes in the Leyden and Berlin Museums. 

Pteropus jJd^^idus, Temminck ; 1825.- — Based on " six individus 
d'&ge et de dimensions diffdrentcs," from the Banda Islands, four 
of which (Temminck writes, I. s. c. p. 186) were kept in the 
Leyden Museum. Of these, three specimens are now in the Leyden 
collection, all mounted, viz. c? ad., skull separate, collected by 
Keinwardt (Cat. ^yst. p. 141, Pt. cjriseus, a; Cat. Ost. p. 255, 
Pt. f/riseiis, a), a very young individual, skull separate, same 
collector (specimen h, skull b), and a 5 ad., skull in sitv., collected 
by Forster (specimen c). Skull a is the original of Temminck'a 
figures, pi. xv. figs. 8, 9. — Two specimens in the Berlin Museum, 
2632 ( (5 ad.) and 4755 ( $ ad.), mounted, skulls separate, have the 
original hand-written Leyden label, with the words " Pteropua 
paUidus, Temra. Mon. ; Banda; Reis van Keinwardt " ; relabelled 
by Peters (printed label) Pt. r/riseus Geoff., Pt. paUidus Temm., 
with an asterisk after the latter name, indicating them as types. 
They are undoubtedly cotypes of pallidus. Skull 4755 is the 
original of the figures in ' Megachiroptera des Berliner Museums,' 
pi. i. figs. 5, 5rt, 5 6 (" Pt. griseus " on plate). 

Remarks. — In skull and dentition Pt. paUidus is scarcely dis- 
tinguishable from Pt. speciostis, mimt(s, and yriseus. From its 
probably nearest relative, Pt. mimus (Celebes), it differs by the 
colour of the underparts and somewhat smaller size, and from 
Pt. griseus by its much darker colour above and below. 

5. Pteropns griseus, E. Geoff. 
Pteropus griseus (pt.), Dobson, Cat. Chir. B. M. p. 44. 

Pteropus griseus, E. Geoffrey., A nn. Mus. rVHiat. Nat. xv. p. 94, pi. vi. 
(animal) (1810: Timor); Cuvier, P. Anim. i. p. 124, footnote 
(1817) ; Denmarest, Mnmm. i. p. 110, no. 141 (1820: Timor); 
Schiriz, Thierr. i. p. L'Jo (1821 : 'I'inioi'l : Temmimk, Mon. 
ATaiinn. i. Tp. 187, pi. xi. (animal) (]82o: Timor); Lesson, Man. 
Mamm. p. 110, no. 282 (1827: Timor); Dexmnre-^f, Did. ISci. 
Nat. xlvi. p. 364 (1827: Timor); Gr/ii/, Griffith's Anin^. Kimjd. 
V. p. 55, no. 157 (1827 : 'J'imor) ; 7*. Geof rot/, Diet. Class. d'Hii<t. 
Nat. xiv. p. 701 (1828: Timor): /. li. Fischer. Syn. Mavim. 
]>. So, no. 12 (1829 : Timor) ; TJ'ai/kr, Siz-^t. d. Amphib. p. 9 (1830) ; 
LesRon, Hilt. Nrif. Ulamw. (C'cmpL ' Buffon) v. p. 54 (1836 : 
Ti'.iior); T(^mminck, M<»). Mamm. ii. p. 81 (pt.), pi. xxxv. fig. <> 
(hpad). pi. xTxri. fip-s. 12, 13 (skull; nee fig.e. 14, 15) (1837 : 
Snniao. nr. Timor) : Graii. .Mnr/. Zoi^l. S- Rot. ii. p. 503(1838: 


Timor) : Oken, Allg. Naturg. vii. ALtli. ii. p. D90 (1838) ; 
Wagner, Schrebers Siiug., S»ppL i. p. 355 (1889 : Timor and 

iioighbouiing islands) ; S. Miiller, in. Temniinck's Nat. Gesch. 

Nedeii. Overz. Bez., Zoogd. pp.20, 59 (pt.) (1839-44: Timor); 

Lesson, N. Tahl. R.Anhn., Mamm. p. 18, no. 185 (1842 : Timor) ; 

Schinz, Sr/st. Verz. Siiug. i. p. 128 (pt.) (1844: Timor); E. 

JDesmarest, Diet. Unit: d'llist. Nat. xi. p. 249 (1848 : Timor) ; 

Grag, Zool. ' Samarang,' Vert. p. 11 (pt.) (1849: Timor); 

Wagner, Schreber's Siiug., Suppl. v. p. 602 (pt.) (1853-55 : Timor) : 
^ Gervais, Hist. Xat. Mamm. i. p. 187 (pt.) c. iig. (head) (1854: 

Timor); Giebel, Sdn^. p. 997 (1855: Timor); Finsch, Neu- 

Guinm, p. 150 (pt.) (1805 : Timor) ; Meters, MB. Akad. Berliii, 

1867, p. 326 (pt.) (Timor); Fitzinger, SB. Akad. Wien, \x. 

Abth. i. p. 445 (1870 : Timor) ; Marchi, Atti Soc. Ital Sci. Nat. 

XV. p. 516 (1872-73: structure of liairs) ; Dobsoii, Cat. Chir. 

B. M. p. 44 (pt.) (1878: Timor) ; Trouessart, Rev. <§• 3Iag. Zool. 

(3) vi. p. 20b (pt.) (1879: Timor); Seabra, Jorn. Sci. Math. 

Lishoa, (2) V. 110. 18, p. 118 (1897: Timor); Trouessart, Cat. 

Mamm. i. p. 80 (pt.) (1897: Timor); Seabra, Jorn. Sei. Math. 

Lisboa, (2) v. no. 19, p. 167, pi. i. Sg. 4 (palate-ridges) (1898). 
Pteropus (Spectrum) livpomelanus b. griseus, Matschie, Megachir. 

p. 24 (1899); Troiiessart, Cat. Mamm., Suppl. p. 52 (1904: 

Pteropus temminclii [nee Pet.), Trouessart, Her. i^- Mag. Zool. (3) 

vi. p. 205 (pt.) (1879: Timor; Samao) ; Jentink, Cat. Ost. 

Mamm. p. 254, specimen a (1887: Samao); id., Cat. Syst 

Mamm. pp. 140, 141 (pt.) (1888: Samao: Timor) ; Trouessart, 

Cat. Mamm. i. p. 79 (pt.) (1897 : Samao; Timor) ; Willink, Nat. 

Tijd. Nederl. Ind. Ixv. p. 275 (1905: Saniao). 
Pteropus (Sericonycteris) temmincki, 3fatsr/iie, Megachir. p. 31 

(1899: Samao; Timor); Trouessart, Cat. Mamm., Suppl. p. 54 

(1904: Samao). 
' Pteropus livpomelanus {\>t.,nec Temm.), A. B. Meyer, Abh. Mus. 

Dresden, "vii. p. 6 (1899 : Bouerato) ; Willink, Nat. Tijd. Nederl. 

Jho'. Ixv. p. 274 (1905 : Salayer). 

Diagnosis. — Similar to Pt. mimiis, but much paler. Forearm 
1 1 4-5-1 18 mm. I-luh. Timor and Dyampea group. 

Palate-ridc/es. — 5 + 5 + 3 ; no trace of an additional ridge between 
the normal ninth and tenth ridges. Arrangement as in Pt. hgj[>o- 
melanus (antea, p. 101). 

Colour. — The darkest extreme approaches in general aspect the 
palest-coloured individuals of Pt. mimus, hut tbe head, mantle, and 
underparts are considerably lighter and the brownish colour of the 
back more lightened with greyish. The palest extreme comes rather 
near in colour to Pt. temmincl-i. 

Dark extreme: J J'g- ad. al., Bonerato, — Back 
Prout's brown rather thickly mixed with pale greyish, producing 
the general effect of an indeiinite greyish brown. Sides of back, 
along membranes, and forearms pale buffy tinged with wood-brown ; 
rump and thighs similar, but tinged with pale cinnamon. — Breast, 
belly, flanks, and anal region pale buffy, rather yellower (more 
golden in tinge) on breast and flanks than on belly and anal 

rrr:i:orcs gkisk;s. J3!? 

region. Cjricealed b.ises of hairs everywhere biowuisl). — Muntlo 
buft'y, titrougly contrasting with back, palest (cream-bufFy) pos- 
teriorly on shoulders, tinged with tawny orange-buff on nape of 
neck and shading into a deeper tawny on sides of neck and foreuetk. 
Short bases of hairs of mantle and sides of neck dark brown ; hairs 
of forcneck uniform to extreme base. — Crown pale cream-buff with 
concealed brownish bases to the hairs, gradually shading on occiput 
into tlic colour of the mantle. Sides of head similar to crown but 
Bomewhat blotched with tawny. Throat mixed pale greyish and 
dark brown. 

Intermediate colour phase : 5 skin, quite or almost full-grown, 
Dv'ampea, — Uack greyish sprinkled with brownish hairs, the 
former colour predominant, producing the general effect of a tinge 
approaching smoke-grey. — Ereast, belly, Hanks, and anal region 
light wood-browu with a delicate tinge of pale cinnamon, more 
distinct on Hanks and anal region than on breast and belly. Con- 
cealed bases of hairs dark brown.— Mantle pale cinnamon wood- 
brown, shading into cream-huffy posteriorly, and into a slightly 
deeper cinnamon wood-brown on sides of neck and foreneck. Base 
of hairs of mantle and sides of neck daik brown ; hairs of foreneck 
uniform. — Crown and sides of head whitish grey. Throat similar, 
but somewhat mixed with brownish, particularly in centre. 

Pale extreme: 6 imm. skin, almost full-grown, Bonerato, 
97.1. 3. o. — Back and rump greyish white with n faint tinge of 
cream-buffy particularly on rump ; base of hairs slightly darker. 
Head and underparts aa in foregoing; mantle somewhat ])uler. 

Measurements. On pp. 140, 141. 

Specimens examined. Ten, in the collections of the Paris, Leyden, 
and British Museums, including the type of the species. 

liange. Timor, with Samao ; Bonerato ; Dyampea ; ? Salayer 
(Specimens not examined). 

Type in the Paris Museum. 

Pteropus r/riseus, Geoff'.; 1810. — Based, it would seem, on three 
specimens (" nous en possedons les deux sexes et la femelle, avec 
de longues tetines "), ol)taincd in Timor, during Pcron and Lesueur's 
voyage. Of these one, or perhaps two, arc now in the Paris 
Museum, viz. (1) a female, not quite full-grown, mounted for front 
view, wings spread, skull in skin, colour much faded (Peg. no. A. 42) ; 
probably not the original of Geollroy's plate ; (2) a young skull, 
in the Museum d'anatomie comparc'e (A. 6745), marked ^-Pteropus 
rjriseiis (Geoff.) ; type"; the evidence that this is really the skull of 
one of Peion and Lcsueur's specimens appears to be unsatisfactory. 

Temminck's Pt. r/risens, 1825 (1. s. c.), is Geott'ioy's Pt. f/riseus 
redescribed and refigured by a copy of Geoff'roy's plate. His P(. 
(/riseus, 1837 (1. s. c), is a mixture of two species, viz.: (1) the 
true Pt. griseus; of Temminck's original material (1837) arc now 
in the Leyden iluseum four mounted specimens, from Samao and 
Timor (Cat. Syst. pp. 140, 141, sub Pt. femmiucki, a, b, i, j\ 
and one skeleton, from Samao {Cat. Ost. p. 254, a) ; the skull of 


PTEROPUS gtiiset;!' 

this latter is the original of Temmiack's pi. xxxvi. figs. 12, 13', 
(2) Ft. temrninchi, Pet. (see this species, infm, p. 318). 

llemarl-s. — In skull, dentition, size and shape of ears, distribution 
nnd length of fur, and external size, this species is similar to Ft. 
mimvs and pallidus, from which it differs in the much paler colour 
of the fur. In this latter character it sometimes rather closely 
resembles Ft. temmincki, from which it is at once distinguished by 
the much heavier skull and dentition, naked tibia, and considerably 
larger size. 

n. 5 yg. ad. sk. ; .skull. Dyaiiipea ; end of dry A. Everett [C.]. 

season, Dec. 189."). 
i. 'cJ ; eknll. Bonerato; beginningof A. Everett [C.]. 

rains, 23 Dec. 1895. 
#. C? yg. ad. .il.; skull. Bonerato; Dec. 189.5. A. Everett [C.]. 

External measui-emenis of Pteropus pallidus and griseus. 

Pt. pallidus. 

5 ad. 

(Incl. 2 cotypes.) 

Pt. griseus. 
4ad. * 

Mw. M.\x. 

I mui. mm. 

ForeArm I HSo 119 

Pollex, total length, c. u 

,, metacarpal 

1st phalanx 

2iid digit, metacarpal 58-5 

1st phalanx 14 14 

,, 2nd-3rd phalanx, c. u 14 16 

Si-d digit, metacarpal 79 83 

1st phalanx i 58 62-5 

„ 2nd phalanx j 84 88 _ 

4(h digit, metacarpal I 78 82 5 

1 „ 1st phalanx 1 47 52-5 

I ,, 2nd phalanx ; 47 52 

• {)th digit, metacarpal 81'5 8.5 

t „ 1st phalanx ' 34 38 

■ ., 2nd phalanx ' 34 37'5 

, Earti, length trom orifice 

• ., greatest width, flattened i 

1 Front of eye to tip of muzzle \ 

iLowerleg | ?54-5 

i Foot, 0. u ! 

i Calcar ' 


1 14 o 






























? 50-5 



* B. M. (Boncrafo); Lcydeii Museum, mounted gpecitucn? a, b, 
(S.imrio ; Timor). 



Measurements of shdh and teeth of Pteropus pallidus and griseus. 

rt. pallidus. Pt. grisetis. I 

Skulls: 4 ad. \ Skull*: 1 yg. ad. | 

Teeth: Sad.Jimm. Teeth t: 1,1 

(Incl. 2cotjpes.) 3 imtn. [ 

Skull, total length to gnathion 

„ front of orbit to tip of nasals ... 

„ width of brain-case at zygomata 

„ zygomatic width 

,, width across m' externally 

„ lachrymal width 

,, width across canines, externally. 

,, postorbital constriction 

,, interorbital constriction 

,, between p*-p^, internally 

„ between cingula of canines 

„ orbital diameter 

Mandible, length 

„ coronoid height 

Upper teeth, c-m^ 

Lower teetli, c-io, 

Upper incisors, combined width 

p', length 

„ width 

p'', length 

,, width 

m', length 

„ width 

m^ length 

„ width 

Pi. length 

„ width 

Pa. 'ength 

,, width 

p^, length 

„ width 

ui ,, length 

,, width 

in.„ length 

,, width 

UI3, length 

„ width 











23 8 



































4 5 



MiH. Max. 
mm. mm. 
















2 2 























» B. M. (Bonerato). 

t P.. M. 97.1..117 (Bonerato), (Bonerato,\ (Dyaajpea); 
Levden Museum, skeleton a (San\floi. 

142 PTKROrU!!! SiTYRUS. 

0. Pteropus satyrus, A'. AnJ. 

Pteropus satvrus, A'. Andersen, Ann. S) Mag. N. 11. (8) li. p. 362 
(1 Oct. 1908 : Narcondam). 

Diagnosis. — Similar to Ft. hijpomelanus, but with smaller etye» 
aud longer fur. Blackish above and beneath, with briiihter-coloured 
mantle and centre of breast and belly, i^orearm about 139 mm. 
Hah. Xarcoudam. 

ShuU and teeth. — Skull as in Ft. hi/j:>omelaniii^, but with equal 
general size of skull the orbits are slightly smaller ; diameter 
12-2 mm. against 12-7-13-2 in all races of Ft. hi/pomehtnits. Teeth 
not differing appreciably in structure or si/e from those of Ft. 

Far. — Longer than in Ft. hi/pomelarnis : about 18-19 mm. at 
middle of back. Distribution of fur as in the allied species. 

Colour. — Type ( S ad., leeth almost unworn) : Back and rump 
blackish seal-brown, thinly aud evenly sprinkled with pale greyish 
liairs, producing the general effect of a blackish slightly lightened 
with greyish; greyish admixture rather more cousjncuous on back 
than on rump. — Centre of breast and belly golden buffy, with short 
concealed seal-brown bases to the hairs ; sides of breast and belly, 
anal region, and flanks blackish very slightly sprinkled with pale 
greyish. Blackish sides of breast and belly washed with seal-brown 
and brownish russet before merging into bright colour of centre of 
breast and belly. — Mantle chocolate with deep chestnut bases to the 
hairs, lightening posteriorly, in a transverse line across shoulders, 
to pale chestnut with ornnge-rufous hair-bases, and darkening on 
sides of neck to dark chocolate, this again on forcneck to mixed 
blackish aud chocolate. — Crown and sides of head mixed blackish, 
buffy, and pale greyish ; throat blackish somewhat sprinkled with 
pale greyish. 

A young specimen (not full-grown) has the colour of the back 
similar to that of the type, though less sprinkled with greyish. 
Bright area on centre of breast and belly paler, almost cream-buff. 
Mantle lighter, between cinnamon and russet, with short seal-brown 
bases to the hairs, and darkening to russet on sides of neck and 
foreneck. Crown and sides of head seal-brown somewhat lightened 
with russet; throat blackish. 

Measurements. On ])p. 145, 146. 

Specimens examhied. Two, in the collection of the British 

Ranfje. IS'arcondam, X. Andamans. 
; Type in collection. 

Uemarls. — This species is evidently an Andaman representative 
of Ft. hypomelamis, and probably most nearly related to Ft. h. 
•leminorinn, from the Mergui Archipelago. The general colour of 
ihe back is rather similar to that of an average (jennnorum.. though 
on the whole less grizzled w ith greyish ; the colour of the mantle, 
sides of neck, and foreneck is almost perfectly as in (jaainorum; the 

PTEROPrs FArxrirs. 143 

bead, at least in some specimens (see type), has the same peculiarly 
grizzled blackish and greyish colour ; in Pt. saiyras the centre of 
breast and belly is bright-coloured, while in the ordinary phase of 
geminorum the uiiderparts are usually uniform blackish (-with or 
without conspicuous greyish sprinkling), but specimens occur with a 
distinct trace of bright colour in the centre of breast and belly, so 
that also in this respect there is scarcely more than a difference of 
degree between satyms and fianincrum. The onh' essential differ- 
ential characters of Ft. satyrus, as compared with Pt. h. r/i'minortim, 
would seem to be the conspicuously longer fur, the less amount of 
greyish admixture in the colour of the fur, the constantly brighter- 
coloured centre of breast and belly, and the slightly smaller eyes. — 
In general colour Pt. satyrus comes rather near to the Andaman 
representative of the Pt. melanotus group, viz. Pi. tytleri (p. 227), 
from which it is readily distinguished by its smaller size, much 
smaller skull and teeth, and less developed posterior ledges of pre- 
molars and molars. In the Nicobars it is replaced by a distinct 
species, Pi. faunulus. 

a. cJ ad. sk. ; skull. Narcondam, Anrlu- C. G. Rogers, Esq. [P.]. (i.9.1.1. 

mans ; Oct. 1904. {Type of species ) 

/. ; skull. Narcondam ; Oct. C. G. Ttogera, Esq. [P.]. ( 


7. Pteropus faunulus, Miller. 

Pteropus faunulus, Miller, Proc. TLS. Nat. Mas. xxiv. p. 785 
(28 May, 1902 : Car Nicobar) ; Kloss, Atidamans 8; Nicobars, p. 325 
(1903 : Car Nicobar) ; Trouessurt, Cat. 3Iamm., Siippl. p. 53 
(1904 : Car Nicobar) ; Miller, Fani. ^ Gen. Bats, p. 58 (1907) ; 
Mason, Rec. Ltd. Mus. ii. pt. ii. p. 166 (1908). 

Diar/nosis. — Allied to Pt. hyjjomelanKS. but smaller, with much 
smaller teeth and longer fur. Back brownish, lightening to wood- 
brown on rump; mantle and underparts ochraceous-buft" tinged with 
cinnamon; head similar, but paler and grizzled with greyish. 
Forearm 118 mm. /lah. Xicobars. 

Skull and teeth. — General characters as in Pt. hypomtlanus, but 
size much smaller: total length of skull about 54'5 mm., against 
()l-5-6S-5 in all races of Pt hypomclanus ; m', length4m!n., against 
4-8-5-9. Size and other characters of skull very nearly as in Pt. 
colonus (Solomon Is.), but orbits conspicuously larger, and teeth 

Ears. — Xot differing in shape from those of other typical species 
of the Pt. hypomelanus group. 

Fur. — Length of fur very nearly as in Pt. .<!atyrv.s, longer than in 
Pt. hypomelanus ; longest hairs at middle of back about 10-17 mm. 
Distribution of fur as in the allied s])ecies. 

Colour. — 2 ad. skin, teeth well worn. General colour of back 
dark Front's brown, jiroduccd by mixture of seal-brown and mars- 
brown hairs slightly .«])rinkled with silvery grey. Itump approxi- 
mately M'ood-brown tinged with cinnamon, with long seal-browu 

144 I'lEiaiPL'Ss AO.MItULlTiLUjil. 

liases to the baijs. Colour of back gradually pas$iug inio paler 
tinge of rump.— Underparts, from foreneck to anal region, including 
flanks, ochraceous-buffy strongly tinged witb pale cinnamon, palest 
on flanks, anal region, and foreneck, deepest (approaching pale cin- 
namon) on breast. Base of fur everywhere seal-brown.- — Mantle 
and sides of neck similar to underparte, but rather paler. — Crown 
and sides of head grizzled greyish and buffy -wood-brown, producing 
a paler and more grizzled tinge than that of mantle ; base of hairjj 
seal-V>rown ; throat similar to sides of head, but mixed with blackish. 

Measurements, On pp. 145, 146. 

Specimen examined. One, in the collection of the jBritisU 

Jtange. Nicobar Islands : Car Nicobar, ^ankauri. 

Type in the U.S. National Museum (no. 111730). 

FUropus faunuhts, Miller ; 1902. — Based on skin and skull of 
one male, obtained on Car Nicobar by Dr. W. L. Abbott. The 
measurement of the tibia given by Miller is incorrect ("38 mm.," 
probably a slip for 48). The skin and skull of the British Museum 
specimen have been compared with the type by Dr. Marcus W. 
Lyon, Jr. 

Eemarks.—The complete resemblance in tlie shape of the skull, 
the characters of the teeth, the form and relative size of the ears, 
and the distribution of the fur, are evidence of the close affinities of 
Pt. faunuhts and satyrus with Pi. hypomelaiMs. In the colour of 
the fur Ft. faunulus is very ditferent from Ft. satyrus, but it 
closely approaches F't. hypomelanus enyanns, some eastern races of 
the same species (Pt. 7i. hypomelanus, luieus), and certain small 
species of the Pt. hypomeJanus group, viz. Pt. mimtis and Pt. colomis. 
Both Pt. satyrns and Ft. faunulus are probably offshoots of Ft. 
hypomelanus, but Pt. satyrus would seem to be most nearly related 
to Pt. h. geminorum, from the Mergui Archipelago, while Pt. 
favnultis is apparently more intimately connected with I't. h. 
enf/anus. From this latter form Pt. faunulus differs chiefly in ite 
iBHialler teeth, longer fur, somewhat paler underjiaits, and smaller 

a. $ ad. 8k. ; skull. Nankauri, Kicobars ; C. G. Rogers. Esq. [P.l f,.4.13.1. 
July-Oct. 1W4. 

8. Pteropus admiralitatum, Thos. 

; ' Fteropus admiralitalum, Thotnas, Aim. 8c Mar/. A'. IT. (G) siii. 

p. 293 (March, 1894: Adniiraltv Is.) : Troueesart, Cat.Manim. i. 

p. 82 (1897 : Admiralty Is.) ; 'Miller, Fam. ^- Gen. Bats, p. 58 

Pteropus (Spectrum) admiralitatum (pt.), Blatschie, Megachir. p. 28 

(1899: Admiraltv Is.); Trotte>:mrt. Cat. Matnm.. Sirppl. p. 5-j 
" ■ (1904: Admiralty Is.). 

Diagnosis. — Allied to /'/. liypomelanus, but with smaller ear.s, ttnd 
longer and differently coloured fur; uxiderparts seal-brown heavily 



sprinkled -wilh butl'y grey. Forearm 11S-12G mm. ]]ah. 
Admiralty Is. 

Skull and teeth. — General characters of skull and teeth as in 
Ft. hypomehinus. Size of skull noticeably smaller, nearly exactly 
as in Ft. mimvs. Size of teeth as in Ft. mimus or the smallest- 
toothed individuals of Ft. Jt^/pomelami.?. 

Ears. — llelatively smaller than in the t5'pical races of Ft. Jnfpo- 
melanus, but not differing in shape ; length of ear from orifice 
21-22 mm., against 25-5-27 mm. in Ft. hypoindanns lateus; width 
of ear 15-16, against lS-20. 

Fur. — Longer than in Ft. 7ii/pomelam(ti ; approximate length at 
middle of back 16-18 mm., against 10-14 in the allied species. 
Distribution of fur as in Ft. hypomelanus. 

Colour. — Four specimens (two skins, two alcoliolic), teeth some- 
what \^orn, including type. Back and rump dark Front's hro\\-n 
a})proiiching seal-brown thickly mixed with shining silvery grey 
hairs, most; of these latter with a peculiar tinge of buff or olive- 

External measurements of Pteropus satyrus, faunulus, and 


I'ollex, total length, c. ii 

„ metacarpal 

,, 1st phalaus 

2ud digit, metacarpal 

,, 1st plialans 

,, 2ud-3rd pbalanx, c. 
.■Jrd digit, metacarpal 

„ Isl pbalaux 

„ 2nd phalanx 

-llii digit, metacarpal 

„ 1st phalanx 

„ 2nd phalanx 

5tli digit, metacarpal 

„ 1st phalanx 

2iid phalanx 

Ears, length IVom orifice 

,. gre<itest width, tlattened 

Inter lenioru 1 

Lower leg 

Foot, c. u 




I'i. j 



admiralitatum. . 

6 "d. 

2 ad. 

4 ad. 1 


(Incl. type.) j 

■ -■'■. 

Mrs. Max. 



mm. mm. 



118 126 



50 53 



11-5 12 



25 28 



60 63-5 



12 14 



13 15 



79 84 



565 005 



86-5 88-5 



75-5 81 



465 50 


47 51 



&'i 86-5 ; 



.34 37 



33 .'i6 

21 22 '■ 

15 n; 



51 57 j 


;'.r,-5 38-5 1 


13 5* 

U 15-5 

Approximate measurement (skin). 



Measurements <[f sJcidh and teeth of Pteropus satyi'us, 
faunulus, and admiralitatum. 

Ft. sati/ncs. 




I Ft. faunulus. 
I '2 ad. 

j Ft. admiral- 
\ itatum. 
j Skulls & teeth : 
I 4 ad. 

I (Incl. type.) 

Skull, total length to gnathion 

„ palation to incisive Ibramina 
„ front of orbit to tip of nasals 
„ width of brain-case at z_vgoniata 

,, zygomatic width 

,, width across m\ externally 

„ lachi-ynial width ." 

„ width across canines, externally, 

„ postorbital constriction 

,, interorbital constriction 

„ width of mesopterygoid fossa ... 

between p*-p', internally 

,, between ciijgula of canines 

,, orbital diameter 

Mandible, length 

,, coronoid height 

Upper teeth, c-m^ 

Lower teeth, c-nig 

Upper incisors, combined width 

p3, length 

„ width 

p^ length 

,, width 

m', length 

„ width 

m'-, length 

,, width 

Pi, length 

,, width 

P3, length 

„ width 

Pj, length 

,, width 

nil, length 

,, width 

m.^, length 

„ width 

nij, length 

,, width 





















^ i 

2-4 ' 

2-1 i 

2 I 

1-8 ' 

4-2 ■ 

2-8 1 



4-8 ! 

2-8 I 

3-9 i 


2-2 j 


(54-5 1) 

































































B. M. ( 

t Estimate. 

I'TKKOl'CS roi.O.Ni ■). 147 

buH', iiroJiiLUig the total effect of a liair-browu or ulive hair-brow ii. 
— Underjjarts from chin to crissiiiu, except a more or less distinct 
narrow collar across tbreneck, dark seal-brovvu thickly sprinkled with 
glossy whitish-grey or buify-grej- hairs ; according to the greater 
or less admixture of greyish, the general colour is whitish-grey 
or buffy-grey mixed with seal-brown, or seal-brown heavily mixed 
with whitish-grey or buffy-grey. — Mantle ochraceous-buff (lightest 
extreme) or approximately ochraceous, or pale hazel tinged witlx 
ochraceous, the colour gradually darkening on sides of neck, and 
continued across foreneck as a narrow collar more or less obscured 
by admixture of the general colour of the underparts. Base of hairs 
of mantle and sides of neck seal-brown. — Crown and occiput light 
buffy hair-brown, becoming gradually darker on sides of head, and 
shading on throat into the general colour of the underparts. In 
one specimen, which on the whole represents the lightest extreme 
in the small series, the crown and occiput are nearly cream-buff, 
gradually darkening on sides of head, and shading on throat iuto 
the general colour of the underparts. 

Measurements. On jip. 145, 146. 

Specimens examined. The Eritish iluseum material. 

Uanye. Admiralty Is. 

Type in collection. 

Remnrlcs. — This species probably replaces Ft. Itifpomelanns in I he 
Admiralty Islands. In the colour of the fur it approaches the 
North Polynesian species of the Ft. mariannus type. 

a.; skull. Admiralty Is. Lords of tlie Treasury 

vCliallenger Exp. i. [P ]• ( 7.'w '^f species ) 

b. (;? ad. sk. ; skull. Admiralty Is. Lords of the Treasury 80.1L2'l.o. 

(Challenger Kxp.). [P.]. 

erf. Jad.,; Admiralty Is. ; Lords of the Treasury !I0.2.20.2, 3. 

skulls7 March, 1875(Oial- [P.]. 
lenger Exp.). 

9. Pteropus colonus. A'. Ami. 

Pteropus hvpomelanus (nee Tenim.), Thomas, P. Z. S. 1887, p. .^■"22 

(Alu, Shortland L) ; id., P. Z. S. 1 888, p. 47 1 ; Trouessart. Cat. 

Mnmm. i. p. 82 (pt.) (1897 : Solomon Is.). 
Pteropus (Spectrum) hvpomelanus (pt.), Matschic, Megaehir. p. 24 

(1899: Alu). 
Pteropus colonus. A'. .4v(lersen, Ann. ^- Mag. K. H. (8) ii. p. 3ii.3 

(1 Oct. 1908: Shortland I.). 

Diagnosis. — Similar to Ft. hi/pomelanus Ixteits, but mucli smaller, 
with shorter ears, and much darker Vireast and belly. Forearm 
ll»9-5-114 mm. JJah. Shortlaid I., W. Solomon Is. 

Skull ttnd teeth. — Essential characters of skull as in Ft. hiipome- 
Ictnvs. but size markedly smaller : total length 55 mm., against 
61-69 in all forms of Pi. hypomelanus ; mandible 4lS-43'b, against 
47*7-54-7 ; orbital diameter 11-2-11-5. against 12- 7-13-2. — 


148 riEKorus solomonis. 

Dentition differing in no important character from that of Pt. liypo- 
melanus, except in the slightlj smaller size of m'. 

Ears. — Size as in Pi. admiralitatum, but tip rather broader and 
more broadly rounded off. 

Fur. — Longest hairs at middle of back about 11-13 mm. Dis- 
tribution of fur as in Pt. liypomelanus. 

Colour. — Type (teeth well worn) and paratype (teeth slightly 
worn). — ^Back and rump Front's brown, rather thinly and incon- 
spicuously sprinkled with greyish-white hairs.- — Breast, belly, and 
Hanks dark Front's brown (type) or mars-brown (paratype), thinly 
(type) or thickly (paratype) sprinkled with grey and greyish-white 
hairs. — Mantle and sides of neck cream-buff slightly washed with 
ochraceous-buff, strongly contrasting with back ; foreneck similar 
to sides of neck, but considerably darkened by admixture of 
brownish. Base of hairs everywhere dark brown. — Crown and 
occiput similar to mantle, passing gradually into a darker tinge ou 
sides of head, and this in turn into the dark brownish colour of the 

Measurements. On pp. 151, 152. 

Specimens examined. Two, type and ])aratype of species. 

Hange. tShortland, West Solomon Islands. 

Type in collection. 

Remarls. — There can be no doubt that tliis species is an eastern 
offshoot of Pt. liypomelanus. In the colour of the fur of the upper- 
side it accords very closely with the New Guinean form of that 
species {Pt. h. hiteus), difiering chiefly in the smaller size, relatively 
shorter ears, and darker underside. In the Central Solomon Islands 
it is replaced by the next following species. 

rt. cJ ad. al. ; skull. Shortlancl I., Solo- Dr. H. B. Gappy 84.3.2,5.1. 

iiions. [C. & P.]. 

b. 5 ad.?li.; skull. Alu, ShorMand I. : C. M. Woodford, Esq. 87.1.18..S. 

April, 1886. [C.]. ( Tf/2^e of species.) 

10. Pteropus solomonis, 7'hos. 

Pteropus solomonis, Thomas, jS'oi\ Zool. xi. p. 597 (Sept. 1904 : 
Ghizo 1.). 

Diagnosis. — Allied to Pt. colonus, but skull slenderer, m' longer, 
fur richer, tibia thickly haired above, colour of fur darker. Head 
and back nearly seal-brown, underparts vandyck-brown thinly 
sprinkled with silvery white, mantle and sides of neck cinnamon 
mars-brown. Forearm 110 mm. Ilab. Ghizo, Central Solomon Is. 

Shid? and teeth. — Skull similar in general size to thac of 
Pt. colonus, but slenderer, rostrum shorter and considerably more 
compressed laterally, mesopterygoid fossa and palate narrower. 
Length of rostrum from front of orbit to tip of nasals 15'5 mm. 
(17-7-1S-2 in Pt. colonus), lachrymal width 11 mm. (12-7), width 
of mesopterygoid fossa 6"2 mm. (7), width of palate between 

r rr.i>.oi'i\s ukux.v i;r,s-. 


f[>\8 mm (9 9-2). Sagittal crest not fully developed fo„e 
UKhndual ^v.thmuch worn teeth), the temporal ridges beiTg dole y 
approximated but not quite fused posteriorly. -^Teeth scarcely 

lengtli of m : o-2 mm, against 4-4-4-0 i,i Pi. cohnus. 
iiars.— bhape and size as in 7^/. colomts 

thicl'lv-Jf""""" ^"'^- ''t'\ *^'"" ^" ^^'- '°^'''^''^ ^"^ extending 
thicklj aloug proximal three - fourths of upperside of tibia 

tTr^^Sn T^'\°^ ^'^'' ""^'^ ^'5' -^■^tle'^ie, belly 14 mn'" 
Least width of furred area of back about 43 mm 

Culoar-Tyi^e ( $ ad. skin, teeth much worn ; Tv^ovember) 
Back seal-brown with a few silvery whitish hairs. Hump, thi-^hs 
and furred part of tibiae distinctly paler-coloured, owing to^prese^nce 
ofshort pale mars-brown (almost buffy mars-brown) tips to the 
hairs.--Lntire underside vaudyck-brown, sparsely sprinkled with 
some silvery whitish hairs on breast and belly, dkenin. to seal 
brown along centre of breast, and becoming slightly paler almost 
walnut-brown on foreneck.-Mantle and ''sides^f^S'b twee. 
cmnamou and mars-brown, not very strongly contrasting wit 
foTb^ rir "/ T'^' «eal. brown for basal half, on sides o1 neck 
similar ohi~^;"r^? T'"''-' "^t'^™^="l'^'- ^P^^e, and sides of face 
Z- . , f 1 '' /,! "^^'^^ '^'''''^ ^^'^ g^^yi-^1^ ^""^ buffy russet, 
TTrfK to V V P,?''^'' "^ paler-coloured tips to the hairs 
partlj to a shght S])rinkhng with silvery whitish hairs. Tempora 
region similar in colour to chin and throat. ^ 

Measurenients. On pp. 151, 152. 

licuuje. Ghi/.o Island, ^ew Georgia group. Central Solomon Islands 
V^;;<' in collection. 

Mwtory in Uterature.~l^hn^ far (December, 190S) the type is the 
only si)ecimen recorded in literature. ^^ 

thf7rj''~'^^y '''^"""'"'' '' *'''' ''^*^^^'°« ^^«t^™ representative of 
rclati>; ^%^'Z '^'<^^n]- F^m its probably nearest living 

rS w f,""?-^^^- ^"^°'"°" ^^-^'it differs in the characters 
referred to in the diagnosis and description above. Prom Ft 1.1- 

dish-;/ ;• ^""'^'f l^"'^^' ^^'"^J^ it approaches rather closely in the 
dist bution and colour of the fur, it is readily distinguished by i ^ 
sma , uehcately built skull, much weuker'dentition, r ative y 
smaller eai's, and somewhat smaller size. • 

n. 2 afl 

kliir"'' ^'';^";.. S°''^""^"I«-: A^"v. 10, A. S. Meek [C]. 
^ -^- {T//pe of species. } 

1 1 . Pteropus brunneus, Dubson. 
Pte.-oims bnuDieus, Dobsou, Cat. Chir. B. :Nr. p. ;J7. 

^1v?ov'n'''""r'''' ^'^'°"l ^- '; '■ P^- "'• %■ * '■^^'^'•> (-^""e, 187S : 
i cu> I.) ; Oy,//;,/, Caf. Austral. M„mm. n. 7!) ( JSit^ ■ Percv I ) • 


coast opposite Percy I.) ; Trouessart, Cat. Manim. i. p. 79 (1897 : 
Percy J.) : Miiler', Fam. S,- Gen. Bats, p. 58 (1907). 
Pteropiis (Spectrum) brunneus, Mutschie, Megnchir. p. 22(1899); 
Trouessart, Cat. Mamm., Suppl. p. ol (1904: Percy 1.^ 

Diagyiosis. — Skull nnd dentition typical Pteropine. Ears 
moderate, exposed. Tibia clothed above. Fur above and beneath 
dark brown (approximately Prout's brown) varied with paler tips 
to the hairs, mantle russet. Forearm 118 mm. Uah. Percy Island. 

SIcull and deniiiion. — Skull in every respect similar to that of 
Pt. Mjpomelanus. Dentition as in the large-toothed races of that 
species (^Ft. h. canus and lejndus). 

Ears (described from one skin). — Moderate, exposed. Inner 
margin evenly convex from base to tip ; outer margin flatly convex 
in basal half, almost straight ia apical half; tip narrowly rounded 
off. Basal half of conch clothed with short, very thinly scattered 
hairs behind, terminal half naked. 

Interfemoral. — Scarcely detectable in centre. 

Fur. — Somewhat adpressed on hack, spreading on rump. Length 
of fur moderate: at middle of back about 16, middle of mantle 17, 
middle of belly 13 mm. Least width cf furred space across back 
about 45 mm. 

Above, proximal third of forearm with short, thinly spread, 
adpressed hair. Tibia clothed, the hair being long and dense on 
proximal three-fourths, gradually shorter and more scattered on 
terminal fourth. — Beneath, short dense woolly hair on antebrachial 
membrane, along outer side of forearm, and on lateral membrane 
between humerus and femur. Proximal third or half of tibia 
liairy, distal portion practically naked. 

Colour. — Type, J ad. skin, Percy I. ; teeth slightly worn ; 
perhaps a little faded. Back and rump approximately Prout's 
brown, paler laterally in a narrow line along membrane. Individual 
hairs almost seal-brown at base, ochraceotis-buffy at tip, the dark 
colour everywhere, except along membrane, showing through : 
hence the approximately Prout's brown total impression of the 
colour. — Entire underside (chin, throat, breast, belly, and flanks) 
paler than back, between Prout's brown and mars-brown, shading 
to vandyck-brown along middle of breast and belly, and almost 
evervwhore sprinkled with some glossy huffy hairs. — Mantle russet, 
becoming paler (nearly clay) on shoulders, and shadiag to dark 
russet OM sides of neck. Concealed base of fur of mantle and sides 
of neck seal-brown. A tuft of huffy glandular hairs on each side 
of nook, nearly covered by surrounding dark fur (probably sexual 
character). — Crown and sides of head mixed Prout's brown and butfy 
russet, all the hairs being seal-browu at base, with ochraceous 
or golden ochraceous tips. 

Measurements. On pp. 151, 152. 

Specimen examined. One, the type. 

Range. Percy I., off E. Queensland, Australia ; ? adjacent coast 
of Australian continent (" a camp of Pt. brunnms " is stated to have 



been observed, by Mr. Broadbent, flj'iug over the Australian coast 
opposite Percy 1. ; see Lucas, I. s. c). 

Type in collection. 

Hidory in literature. — jNTothing has been added to our knowledge 
of this species, since Dobson described it, in 1878. 

liemarks. — The perfectly typical, hf/pnmelanus-like skull and 
dentition of this species leave no doubt that it is an Australian 
representative of Ft. hypomelanus. Erom the geographically nearest 
form of that species \Ft. h. luteus, New Guinea) it is readily 
distinguished by its furred tibiae, much darker colour, and smaller 
size. Externally (colour, size) it bears no small resemblance to 
l^t. cof/nalus (San Christoval, Solomon Is.), which however differs 
essentially in skull and dentition. 

J ad. sk. ; 

Percy I., E. Queens- 

Voyage H.M.S. ' Herald ' 



(C'apt. Deiibam). 

( Tt/pe of species.) 

E.vternal measurements of Pteropus colonus, solomonis, and brunneus. 

Vt. colonus. 

2 ad. 

Type and 


Ft. solo- 

Pt. hrun- 

2 ad. 

6 ad. 

2 ad. 
I Type. 

j nun. 

Forearm j luyij 

Pollex, total length, c. u | 43'5 

„ metacarpal 11 

„ 1 st phalanx | -!4 

2ucl digit, metacarpal 60 

„ 1st phalanx 11 

,, 2nd-3rd phalanx, e. u. ... 12 

3rd digit, metacarpal | 73 

,, 1st phalanx 510 

,, 2nd phalanx 78 

4th digit, metacarpal I 72 

,, 1st phalanx 1 41 

,, 2nd plialanx 

r)th digit, metacarpal ' 7-!'o 

., 1st phalanx 32 

,, 2nd phi\lanx .. 33 

Ears, length from orifice 

,, greatest width, flattened 


Lower leg 49 

Foot, c. u 

Calcur (12) 


Paratype *. 









2/) .5 





14-5 1 


12 1 


75 ' 





7") 5 















1 18 


* B. M. S4.3.25.i. 


rrEUorrs colonus etc. 

Measurements of sl-uUs awl teeth of rt.erojnis colonus, solomonis, 
CDid bruuncus. 

Skull, total length to gnathion 

„ palation to incisive foramina ... 
,, t'ront of orbit to tip of nasals ... 
,, width of brain-case at zygoniaia 

,, zygomatic width 

„ width across m\ externally 

„ lachrymal width 

,, width across canines, externally. 

,, postorbital constriction 

,, interorbital constriction 

,, width of mesopierygoid fossa ... 

,, between p*-p', internally 

between cingnla. of canines 

,, orbital diameter 

Mandible, length 

,, coronoid height 

Upper teeth, c-m''^ 

Lower teeth, c-ni^ 

Upper incisors, combined width 

p'', length 

,, width 

p^, length 

,, width 

ru^, length . 

,, width 

m-, length 

„ width 

p„ length 

,, width 

Pj. length 

,, width 

Pi, length 

„ width 

mj, length 

,, width 

m^, length 

„ width 

tn,, length 

., width 

PL colonus. 

2 ad. 
Type and 
para type. 

$ ad. c? ad. 
Type. Paratype. 










23' 2 






















7 ■ 
















Pt. solo- ' Pt. hrun- 
'DW^iis. I neus. 

2 ad. 

c? ad. 














2 2 

































TTBKOrr.S ORXAXr.S; 153 

I'J. Pteropus ornatus, Gra;/. 
Ptfii-opus vetulus (pt.), Dobson, Cat. Chir. B. M. p. 27. 

Pteropus rubricollis (nee E. Geoff.), Joiian, Mem. Soc. Imp. Sci Sid. 
Cherbourg, ix. p. 89 (18(i3 : >»e\v Caledonia; luibits) ; id., op. c. x. 
p. 301 (1864 : New Caledonia). 

Pteropus vetulus' (nee Joikui), Peters, MB. Akad. Berlin, 18fi7, 
p. 8L>3 (New Caledonia); Dobson, Cat. Chir. B. M. p. L'7 (pt.) 
(1878 : New Caledonia) ; id., P. Z. S. 1878, p. 874 (pt.) (1879 : 
New Caledonia); Trouessart, Rev. !<,• Mag. Zool. (3) vi. p. 204 
(pt.) (1879 : New Caledonia) ; id., Ann. ISci. Nat. (tJ) ZooL viii. 
Art. 12, pp. 16, 17 (1879 : remarks on distr.) ; Jentink, Cat. 
Ost. Mamm. p. 253 (1887 : New Caledonia) ; id.. Cut. Syst. 
Mnmm. p. 139 (1888 : New Caledonia) ; Trouessurt, Cat. Mamm. 
i. p. 78 (1897 : New Caledonia) ; Seabra, J. Sci. Math. Lisboa, 
(2 ) V. no. 19, p. 166 (1898 : New (Jaledonia) ; Matschie, Mer/ac/iir. 
pi. i. iigs. 2, 2 rt, 2 6 (skull) (1899 : New Caledoniii) : Etheridije, 
Proc. Linn. Soc. N. S. Wales, xxiv. pp. 274, 275 (1899 : fur used 
for the spear-becket of the Caledonians). 

Pteropus (Spectrum) vetulus, Matschie, Mec/ackir. p. 21 (pt.) 
(1899 : New Caledonia") ; Troiicssart, Cat. Mamm., Sitppl. p. 51 
(pt.) (1904 : New Caledonia). 

Pteropus ornatus, ffr«//, Cat. Monh. ^-c. p. 105 (1870: New Cale- 

Diagnosis. — Allied to Pt. lijipomelanus, but with somewhat larger 
teeth, uoticoably heavier corouoid process, longer and less adjjres.sod 
fur extending thickly on upperside of tibia, longer wings and tibia, 
and different colour of fur. Head, back, and underparts dark 
brown sparsely sprinkled with white hairs, mantle huffy white, 
forming a stronglj- contrasting tippet. Forearm about 150 mm. 
Mab. New Caledonia. 

Skull and Uetli. — iSkull similar to that of Pt. hypomelanv.s, but 
rostrum rather more compressed laterall3-, crests stronger, coronoid 
l)roce8s conspicuously heavier and more steeply ascending. — Teeth 
averaging decidedly larger, but scarceh' differing in structure. 

Ears. — Somewhat smaller than in Pt. h>/pomelanvs (but looking 
much smaller, owing to longer fur of head), reaching scarcelj- more 
than half the distance between base of ear and back of eye, when 
laid forward ; basal half or two-thirds couceiiled in the long fur of 
the head. Lower half of inner and outer margins strongly convex ; 
upper half of inner margin nearly straight, of outer margin flatly 
concave ; tip narrowly rounded off. Naked posteriorly, except at 

Win(]s. — Membranes inserted close together on back, separated 
by a space of only about 10 mm. 

Jntirfcmoral. — Extremely short or undeveloi)ed in centre. 

Fur. — liather long, straight, directed posteriorly but not closely 
ndpressed on back and rump. Approximate length of hairs at 
middle of back 20-21, middle ot mantle 23-2-5, middle of belly 
20-21 mm. Fur of back extending laterally on membranes about 
25 mm. beyond their lino of origin from back; least width of 
hairy Hpace acros.- back about 60 mm. 



Above, a thin liuo of rather long, adpressed hairs along humerus. 
Proximal half of forearm (excepting basal portion, at and in 
fi'ont of elbow, which is naked) covered with long, distally 
sliorteniug, thinly spread hairs ; on the external side of the forearm 
the hair can be traced, gradually shortening, to within 45 mm. of 
the carpal extremity. Tibia densely clothed with long hairs, 
extending also in a very thin line along metatarsus to base of 
toes ; a few scattered hairs on upperside of toes. 

Beneath, antebrachial membrane, lateral membrane p-long outer 
side of forearm almost to carpal joint, and between humerus and 
lemur densely clothed with long, woolly, frizzled hairs. Tibia 
naked, except at base along inner margin. 

Colour. — (5 imm. skin,; type of species: — Back 
and rump almost chocolate, somewhat sprinkled with silvery 
greyish-white hairs. Individual hairs seal-brown at base, russet 
at tip, producing the general chocolate-brown effect ; the silvery 
greyish-whifn hairs uniform from base to tip. — Entire under- 
side, including sides of neck and flanks, very dark chestnut, 
slightly but not quite inconspicuously sprinkled with silvery 
greyish-white hairs. Fur everywhere (the uniform silvery greyish- 
white hairs excepted) pale russet or almost cinnamon-rufous at 
base, chestnut or chestnut seal-brown at tip. The general effect 
of the combined colours is darkest (chestnut vandyck-brown or even 
chestnut seal-brown) on throat, foreneck, sides of neck, flanks, and 
anal region, in which the dark hair-tips completely or almost 
completely conceal the brighter base of the fur; palest (chestnut 
russet) on breast and front of belly^, in which the base of the fur 
is partially exposed. — Mantle and occiput cream-buff, almost cream- 
white, forming a well defined tippet, in striking contrast to 
general dark aspect of fur, washed with a shade of buffy russet in 
front, laterally, and behind, near surrounding brown fur. Extreme 
base of hairs of mantle, next to skin, seal-brown. — Crown and sides 
of head chocolate, coarsely mixed with golden ochraceous and silvery 
greynsh-white hairs. Short adpressed hairs on muzzle greyish, 
extending backward above eye as an ill-detined superciliary stripe. 

A second skin (imm., sex indeterminable, is very 
similar to the above, but conspicuously paler (russet) on underside ; 
mantle more pure buff in centre, gradually shading to golden 
ochraceous laterally. 

A third skin ( c^imni.,, in abraded pelage, is consider- 
ably darker on head, back, and rump, owing to abrasion of paler- 
coloured tips of hairs ; mantle whitish in centre. 

Measure»ie»ts. On pp. 157, 15S. 

Specimens examined. The material in the Paris and British 

Range. New Caledonia. 

Type in collection. 

Technical name. — Since Peters (1867, ?. s. e.) identified this 
species with Jouan's Pt. vetulus, it has been referred to in literature 
under that name by all authors except Gray. On the true Pt. 
veiuhis, Jouan, see infra. 

rrrporcs vftulus. 


<i. J imin. sk. ; Pi. de France, New C'ale- ParcLased (Mr. 
skull. donia; Alay 1, 1858. Oumingj. 

{Type uf species.) 
h. c? imm. sk. ; Pt. de France, April Tomes Coil. 7-1.1.252. 

BkuU. 20, 1 858 {H. N. Turner), 

c. Imm. sk. ; skull. New Caledouia (if. iV'. Tomes Coll. 


[Pteropus yetulus, Jounn. 
Pteropus Vetula, Jmian, Mem. Sua. Imp. Sci. JS''at. Cherbourg, ix. p. 90 

(1803: New Caledonia); id., op. c. x. p. '6\)l (18ti4 : New Caledouia). 

Pteropus gerinaini, Bohsun, P. Z. S. 1878, p. 874 (1879: New Caledonia); 

Truucasart, lieu, cj- Mag. Zool. (3) vi. p. 1^04 (1879) ; JJobsun, Rep. Brit. 

Afsoc. IBsU, p. 172; Trouessurt, Cat. Mamm. i. p. 78 (1897;. 

Pteropus (Spectrum) vetulu.s a.germaiiii, Troaessart, Cat. Mamm., Siippl. 

p. 51 (19U4). 

Jouan's two species o/ Pteropus from New Caledonia. — In his Notes on llie 

Fauna of New Caledonia (18(53, I. c.) Capt. Jouan distinguished two species of 

Pteropus, viz. " P(. rubricoUis, Leith. ? " ["Leith." evidently a misprint for 

Lath. ; the correct author's name would be E. Geofl.] and " Pt. Vetula, 

Montrouzier" {Pt. Vetula is probably a manuscript name taken up by Jouan ; 

it does not occur in any of Pere Montrouzier's published papers, so far as 

known to me ; his article on New Caledouia i:i the ' Revue algerienne et 

coloniale,' April-May, 1800, has been looked up for me by Professor Trouessart). 

Probably owing to the lacts that Jouan's description of Pt. vetulus was very 

fragmentary and that one of its principal ditferential characters, as compared 

with his " /Y. rubricoUis," was stated to be its considerably smaller size, Peters 

(MB. Akad. Berlin, 1807, p. 323) regarded " Pt. rubricoliis" as the adult, Pt. 

vetulus as the young of one species, for which he therefore retained the name 

Pt. vetulus, and this view was apparently accepted by later revisers of the 

genus (Dobson, Matschie). But in 1879 Dobson himself described a new 

species from New Caledonia, Pt. germaini, said to differ from Pt. ornatus, 

inter alia, in precisely those two characters which, according to Jouaa, 

distinguish Pt. vetulus from " Pt. rubricoUis," viz., much smaller size and 

longer fur. This fact certainly greatly strengthens the assumption that Jouan 

was right in considering his two species perfectly distinct. So far as the 

available information in literature goes (I have not seen the type of Pt. vetulus 

nor that of Pt. genrMini), the characters and nomenclature of these two New 

Caledonian Pteropi are as follows : — 

The species hesitatingly identified by Jouan with Pt. rubricoUis will (if, as 
here presumed, different from his Pt. Vetula) have to stand as Pt. ornatus, 
Gray, 1870 ; synonym, Pt. vetulus, Peters, 1807, et auclorum plurimorum, iiec 
Jouan, 1863. The only important items in Jouan's description of " Pt. rubri- 
coUis" avs these four : (1) length of bead and body 250 mm.; (2) ears very 
small; (3) fur of head, back, and belly "laineus"; and (4) colour '" roux " 
with mantle " hlanc-jaunatre." All these characters agree vi'xlh Pt. ornatus, 
and the second, third, and fourth absolutely exclude the only other New 
Caledouian species of similar size, Pt.geddiei. — Jouan's second species, " Pt. 
Vetula,'' was stated to diil'er from the foregoing: (1) " par sa taille constam- 
ment plus petite," the length of the head and body benig only 100 mm., and 
(2) by having the fur "plus long, pliltot soyeux que laineux." This species 
wiU have to stand as Pt. vetulus, Juuan. 1803, nee auctorum plurimorum ; 
synonym Pt. germaini, Dobson, 1879. 

Pt. germaini, Dobson. — Summary of original description: — Teeth simple, 
like those of Pi. giganteus. Ears shorter than muzzle, concealed by long fur of 
head. Fur long u,nd woolly, like that of Pt. anetianus ; tibiffi clothed with long 
fur extending to backs of feet. Colour: bend and whole inferior surface of 
body dark blackish brown interspersed with several shiny greyish hairs, 
shoulders and back darker, rump and legs greyer ; fur of mantle pale yellow 
with reddish e.xtremities ; muzzle and superciliaries light greyish brown. 
Measurement.' of type, a not quite adult female, head and body 150 mm., ear 20, 

^M riERorrs adeatus. 

foi-eiirui 120, poUcx 58, thinl digit, metacarpal 76, ]st j)h. 63'."), 2ir1 ph. 89, 
fifth digit, metacarpal 76, Jst ph. ,'58, 2nd ph. 34, tibia 56, foot 43. Resembles, 
i)obson adds, externally to some extent Ft. anetianvs, " but the very ditfereut 
form of the teeth at once distinguishes it" ; from Pt. ornaius, "inhabiting the 
same islands, it is distinguished by the completely difierent colour of the fur 
[Dobson had before him speeimens of Pt. ornatus for comparison], as well as 
by the absence of transverse basal ridges in the molars and premolars." 

Type spechneni'. — The type of Pt. vetidus is nor, in the Paris Museum nor, I 
am informed by Professor L. Corbiere, in the Cherbourg Museum. Dobson's 
statement, in P. Z. S. 1878, p. 874, that the type [in 1878 or 1879] was in the 
Paris Museum is erroneous; there is no evidence in the Registers of that 
Museum that it ever was there ; and his descri))tiou in Cat. Chir. B.M. p. 27, 
of '■ a female (typs of the species) " is taken from a specimen of Pt. ornarus. — 
Pt. gcrmaini was based ovi a single specimen in the Paris Museum. I have 
c."iannned all the specimens in that Museum collected by Germain in New 
Caledonia; none is labelled Pt. gcrmaivi, and none corresponds to the det^irip- 
tion and measurements given by Dobson.] 

13. Pteropus auratus, A'. And. 
Pterojius vetulus (pt.), Pobson, Cat. Chir. B. M. p. 28. 

Pteropus vetulus (pt., nee Jouan), Dohson, I. s. c. (1878: Loyalty Is.) ; 

uL, P. Z. S. Ib78, p. 874 (1879 : Loyalty Is.) ; Trouessart, Rev. 

cV Mai/. Zuol. (3) vi. p. 204 (1879 : Loyalty Is.) ; var., Trouessart, 

Cat. Mamm. i. p. 78 (1897: Loyalty Is.). 
Pteropus (Spectrum) vetulus (pt.), Matschie, Merjachir. p. 21 (1899 : 

Loyalty Is.); var., Trouessart, Cat. Mamm., Suppl. p. 51 (19()4: 

Loyalty Is.). 
Pteropus auratus, K. Andersen, Ann. S,- Mag. X. II. (8) iii. p. 2Z'C> 

(1 Feb. 1909: Lifu, Loyalty Is.). 

Diaijnosis. — Similar to Ft. ornatus iu sliull, dentition (though 
perhaps with slightly smaller cheek-teeth), distribution of fur (tibiii 
clothed above), and general size, but with larger ears and different 
colour of the fur. Head, mantle, and underparts golden ochiaceous, 
back mixed ochraceous-buif and brown. Forearm about 145 mm. 
Bah. Loyalty Is. 

Teeth. — General character.s as in Pt. ornatus. Id the single 
known specimen p"*, m\ Pj, p^, and ra^ are distinctly smaller than 
in three skulls of Pt. ornatus; see measurements p. 158. 

PaJate-ridr/es. — 5 + 6 + 3. First ridge terminating lateralh' at 
front of canines ; second at back of canines ; third between canines 
and p' ; fourth at middle of p'' : iifth at front of p^ ; sixth at 
back of p'' ; seventh at front of m' ; eighth at back of m' ; ninth 
at back of (or closely behind) nr ; tenth and eleventh behind nr ; 
twelfth to fourteenth situated at palation border. 

Ears. — Conspicuously longer and broader than in Pt. ornaius ; 
length from base of orifice 24'5 mm., against 20 in the allied 
species, greatest breadth (Hattened) 17 mm., against 12. 

C'oZowr (type). — Back vandyck-brown with long golden ochraoeous- 
buff tips to the hairs, producing the general effect of golden 
ochraceous clouded with brown. — Breast and belly rich golden 
ochraceous, palest (golden ochraceous-buff) at base of hairs, shading 
to tawny on foreneck. sides of neck, and flanks, and to tawny 
nisset faintly sprinkled with ochraceous on chin, throat, and anal 
legion. — Mantle rich golden ocliraceous-buff, this colour confined to 
lips of hairs, middle portion of individual h'lirs buff, extreme base 



mxt to sk.n seal-brown ; colour of mantle shaclin- ffraduallv into 
tawny onocc.putandsidesof neck.-Crown bultV, sTio-htiy darkened 
w,t , brownish, passing- gradually into tawny on^Vides of head ' 

Measurements. Below and on p. 158. 

Specimen examined. One, the type. 

Ramje. Loyalty Islands (Lifu). 

Type in collection. 

Histon, in Uterature.-iiomQ of the differential characters of this 
species were already pointed ont by Dobson (1878 ] s e ) Jho 
however, considered it only a local variety of Ft^ vehdus ml 
statement that the ears are shorter than in the .4; c' edonSn 
species IS erroneous. '-'aicuoiiian 

colour ofST^^T^^- ''""^i"^.'^' ^^''''^' ^" ^'""''^^ «^P^«t, the 
colour of P^. aumtu^ is easily derived from that of its closest rela- 
tive i^.o..«^«. The dark brown of the head and underparts of 
Ft ornatus is in Pt. auratvs replaced by golden ochraceous or 
ochraceous-bnff, and the dark broin of th'brack by mfxed ' den 
ochraceous-buti and brownish. ^omtw 

2 ^^- al. ; skull. Lifu I., Loyalty Is 

Kev. S. J. Whitinee 77.7.:2.S.I. 

[C.]. (TV-/'*-' of species.) 

External measurements o/ Pteropus ornatus and auratus. 

I Forcai'm 

Pollex, total length, c. u. ......... 

,, iiietafarpal 

I „ 1st phalanx 

2iid digit, metacarpal .'. 

., 1st phalanx 

j _ .. I'nd-ord phalanx, c. u. 

I 3rd digit, raetacarpal 

„ Lst phalanx 

I „ 2nd phalaux 

: 4th digit, metacarpal ..", 

[ II 1st phalanx 

I, "Jiid phalanx 

5tli digit, metacarpal 

„ 1st phalanx 

I. 2nd phalanx 

Ears, length from orifice 

„ greatest width, flattened .... 

Front of eye to tip of muzzle 

Lower leg 

Foot, c. u 


Pt. ornatus 

Pt. aziratus. 


9 ad. 

Type. 1 


< mm. 





! 14-2 












J 07 





















■.^ * J'^'^^^ ^''"*' '^'''"' Caledonia, Paris Museum ■ measured hv V,.r,f^ 
])r. E. Trm.esBart. It should be noted tlu.t this indiridual rrle. Is pr£lv 
Tery nearly the maximum size of the species ("c'est le 01.^—1^^'''''' 
que nous possedious dans lalcool." ^L.^savt. irm")^%X'£jT;r^ 
Mu.«euni .«iHHMn.ens are blightlv immature -^"^ inue Uutish 

15S riKROPLS OilXATUS A\U iLKlTl'3. 

Measurements of s /mil of Pteropus auralus, and teeth of Ft. ornatus 
and auratus. 

Pi. ornalus. Ft. auratus. 

Teeth ; 3 iniiu. J ad. 

,, , MiN. Max. | 

tnin. uiui. mm. 

Skull, totallengtU to gnathion (;7-3 

„ palation to incisive fbraiuiria 34 

,, front of orbit to tip of nasals 22'2 

. , „ width of brain-case at zygoinatu i 23-8 

I., „ xygouiatic width i 37'2 

I ,„ width across m\ externailv ■ 188 

r ' ,, lachrymal width 14-2 

j „ width across canines, exterually 12-8 

I I „ postorbital constriction .1, , , j,.i,,. fi'T 

i „ inl.erorbital coristriction ! ' 8-5 

I „ width of iiiesopterygoid fossa 7-7 

' „ between p-*-p^, internally l()-6 

,, between cingiila of canines 7 

orbital diameter 12-8 

I Mandible, length ,54-3 

i ,, coronoid height 29 

I Upper teeth, c-m''' 26-2 

i Lower teeth, c-mj 29-7 

j Upper incisors, combined width 6"9 7'5 7 

|p\Iength 4-fj .5-1 47 

I .. width 3-2 3-3 3 

p*, length 1 48 57 5 

i ,■ width 3-7 3-9 30 

m'. length (J fi-S 5-7 

. n width 3-2 3-7 I 3-2 

jm^, length 28 3-2 ' 2-8 

I M width 2-2 2-3 1-8 

Pi, length I 2-1 2-8 2 

width ,..j 2 2-2 2 

P3. length I 5 5.7 48 

- width 2-8 3 2-5 

P4. length ' .5-2 TvS 5 

.. width 2-9 3-5 28 

nij, length : 5-2 62 5 

,, width ... 29 3-2 28 

m^. length | 4.0 4-8 4I 

., width , 2-7 3 28 

ajg, length I 2'8 3 27 

„ width j 2-1 2-2 ; 21 


14. Pteropus dasymallus, Temm. 
I'ternjnis dasymallus, Dobson, Cat. Chir. B. M. p. 25. 

Pteropus rubricollis {nee I!. Geoff.), G. T. [misprint for Ph.F.'\ Siebold, 
De Hist. Nat. in Japoniu statu, p, 13 (]824 : Nagasaki ; Jeddo). 

Pteropus dasymallus, Tetmuinck, Mon. Mamm. i. p. 180, pi. x. 
(auimal), p'l. xv. fi^s. 10, 11 (skull of type) (1825: Nagasaki j 
Jeddo) ; IJesmarest, Diet. Sci. Nat. xlvi. p. 861 (1827 : Nagasaki ; 
Jeddo) ; Is. Geoff roy, Diet. Class. cVHist. Nat. xiv. p. 700 (1828 : 
Japan) ; J. B. Fischer, Syn. Mamm. p. 83, no. 6 (1829 : Jeddo ; 
Nagasaki); Wayler, Syst. Amph. p. 9 (1830); Lesson, Hist. 
Nat. Muinm. {Compl. Biffon) v. p. 59 (1836; Japan); Gray, 
May. Zool. ^- Bot. ii. p. 5(33 (1838 : Japan); Olen, Ally. Naiurq. 
Yii. Abth. ii. p. 990 (1838); Temminck, Tijd. Nat. Gesch. V. 
pt. 4, p. 282 (1839 : Japan) ; Wayner, Schreber's Sauy., Suppl. i. 
p. 349 (1839: Nagasaki; Jeddo); Temminck, Fauna Japon., 
Mamm. p. 12 (1842 : S. Kiushiu ; Yakushima) ; Lesson, N. 
Tahl. Reyne An., Mamm. p. 13, no. 176 (1842 ; Japan) ; Schinz, 
Syst. I'ierx. »Saw^. i. p. 124 (1844 : Japan); trf.,o/).cj7.ii. Naelitiag, 
p. 14 (1845 : S. Kiusbiu ; Yakushima) ; Gray, Zool. ' Samarany,' 
Vert. p. 34 (1849 : Japan) ; Wayjier, Schreber's Sliuy., Supj)l. v. 
p. 598 (1853-55 : Japan) ; Gervais, Hist. Nat. Mamm. i. p. 188 
(1854: Japan); Giebel, Sauy. p. 997 (1855: Japan); Schleycl, 
Dierkunde, i. p. 53 (1857 : Japan) ; Gerrard, Cat. Bones Mamm. 
B, M. p. 56 (1862 : Japan) ; Peters, MB. Akad. Berlin, 1867, 
p. 323 (Kiusbiu; Yakushima); Fitzinger, SB. Akad. Wien,\x. 
Abth. i. p. 428 (1870: Jeddo; Nagasaki); ScJdeyel, Dierentuin 
Nat. Art. May., Mamm. p. 66 (1872 : Japan) ; Dobson, Mon. Asiat. 
Chir. p. 16 (1876 : Jfipan ) ; id., Cat. Chir. B. M. p. 25, pi. iii. fig. 1 
(ear) (1878 : Japan) ; Trouessart, Rev. ^- May. Zool. (3) vi. p. 204 
(1879: Kiushiu; Hondo); Jentink, Cat. Ost. Mamm. p. 252 
(1887: Japan) ; id.. Cat. Syst. Mamm. p. 138 (1888: Japan) ; 
Fhma7in, Mitth. D. Ges. Naturk. Ostasiens, \. p. 389 (1892 : Liu- 
tius) ; Trouessart, Cat. Mamin. i. p. 77 (1897 : Lin-kius) ; 
Matschie, Meyachir. pi. i. figs. 1, \a,\b (skull) (1899); Banys, 
Amer. Naturalist, xxxv. no. 415, p. 561 (1901 : Isbigaki) ; 
Bonhote, Nov. Zool. ix. p. 628 (1902: Liu-kius) ; Miller, Fam. ^ 
Gen. Bats, p. 58 (1907). 

Spectrum dasymallum, Gray, Cat. Monk. ^-c. p. 101 (1870: Japan). 

Pteropus rubricollis (dasymallus), Seitz, Mitth. D. Ges. Naturk. 
Ostasiens, v. p. 363 (1892 : Kiushiu). 

Pteropus (Spectrum) dasymallus, Matschie, Meyachir. p. 27 (1899) ; 
Trouessart, Cat. Mamm., Suppl. p. 53 (1904 : Liu-kius). 

Diagnosis. — Skull and dentition unmodified Pteropine. Ears 
rather small, half exposed ; fur unu.sually long and spreading ; 
tibia clothed above. Head, back, and underparts dark brown, more 
or less varied with bufFy hair-tips on back ; collar buffy or cream- 
bufiy. Forearm 125-5-137 mm. Bab. S. Liu-kiu Islands. 

Skull and teeth. — As in Pt. hyjiomelanus, but rostrum slightly 
shorter than in specimens of hypomelajius of similar size, p' and p^ 
a little heavier. Compare measurements of skulls and teeth, p. 171 , 
with those of Pt. hypomelanus, p. 131 . 

Pa/a^€-r(V(7Cj> (several specimens). — S-fo-^-S; the right aiid left 


half of tbo sixth ridge ouly slightly separated, in some indivi- 
duals completely fused in the median line (formula 6-f4 + 3). 
First ridge terminating laterally at front of canine ; second at back 
of canine or between this and p' ; third at front of p^ ; fourth at back 
of p'' or between p^ and p^ ; tilth at p^ ; sixth at front of m' ; seventh 
at back of m^ ; eighth at m" ; ninth closely behind m^ ; tenth far 
behind m'' ; eleventh to thirteenth near palation border. 

Ears. — Rather smaller than in J't. hyponulanus (but looking 
much smaller, owing to basal half of conch being hidden in the 
long fur) ; not nearly reaching hinder corner of eye "«"hen laid 
forward. Inner margin of conch flatly convex from base to 
tip ; lower half of outer margin strongly convex, upper half 
ajjproximately straight, becoming very flatly concave immediately 
below tip ; tip subacutely rounded, liasal lialf thinly haired behind, 
concealed by the fur of the head ; distal half naked, exposed. 

Interfemorul. — Undeveloped in centre. 

Fur. — Long, dense, spreading, extremely soft and silky. Ap- 
proximate length of longest hairs at middle of back '2-i, middle of 
mantle 25, middle of belly 20 mxii. 

Above, fur extending on proximal half or two-thirds of forearm, 
and on lateral membrane almost to a line between elbow and knee. 
Tibia clothed to ankle, the hair being long and dense on proximal 
two-thirds or three-fourths, shorter and more thinly spread on distal 
third or fourth. A few long (blackish or whitish) hairs on feet. 
Interfemoral completely covered along femur and proximal half of 
tibia, practically naked laterally along distal half of tibia. — Beneath, 
basal half of forearm hairy. Antebrachial membrane, lateral 
membrane along outer side of forearm, and along body between 
humerus and femur, covered with densely set woolly hair. Proximal 
half or third of tibia hairy. 

Colour. — (S ad. skin, Kobama, S. Liu-kius, July 20th, teeth 
worn ; B. M. 5. 11.''). 13 : Back and rump nearly seal-brown, strongly 
varied with buffy tips to the hairs. Individual hairs seal-brown 
for basal five-sixths or three-fourths, buffy or golden ochraceous- 
buflfy at extreme tip, the pale tips becoming longer posteriorly, 
on rump and thighs. lu most places the pale hair-tips are too 
short to completely cover the seal-brown bases ; many of the hairs, 
particularly on the sides of the back and distal two-thirds of tibia, 
are seal-brown from base t« tip. — Breast, belly, and flanks much 
more uniform in colour : dark vandyck-brown, approaching seal- 
])rown, in some places (especially on flanks) washed with mars-brown. 
I'oncealed base of hair everywhere almost seal-bi'own. A few 
silvery greyish- white hairs interspersed between the dark ones. — 
Mantle and sides of neck buffy (approximately buff-yellow), sliading 
to whitish cream-buff on sides of forcneck. Pale collar iiiterrupfed 
in median ])ortiou of foreneck by a narrow line of daik brown 
connecting the dark throat and breast. Concealed basal half of 
hairs in central portion of mantle dark brown ; at sides of mantle, 
sides of neck and foreneck the dark brown is confined to the 
extreme base of tlie hairs or completely obliterated. — Occiput, 

PTKROPUs nAsv.UAi,r,r.s. Ifil 

crown, iiitorooular region, and sides of liead dark brown (apiiroachiiij; 
seal-brown), slightly grizzled with pale grcj-ish and buffv ; chin and 
throat almost uniform seal-brown. 

Individual variation (skins, B. Liu-kius) : — Not great, cliiefly 
depending on the lesser or greater development of buffy tips to tlic 
hairs of the back and rump, the more seal-brown or more mars- 
brown tinge of the ground-colour of the fur, the amount of grizzling 
on the head, and the richer or paler colour of the mautle. — In soino 
specimens the buffy hair-tips ou back and rump are extremely short, 
and almost ochraceous iu tinge, the general colour of the back and 
rurap therefore seal-brown slightly varied with dull ochraceous. 
In others, the buffy tips are so long as to almost completely cover 
the dark base of the fur, making the back and rump approximately 
pale buffy, with a narrow line of mars-brown along the lateral mem- 
brane (9 ad., Ishigaki, June; — Breast, belly, and flanks 
varying from seal-brown tinged with vaudyck-brown to almost 
uniform pale mars-brown. — ^ilantle varying from almost buff-yellow, 
through pale ochraccous-buff, to pale butf ; side of foreucck always 
much more whitish in tinge than mantle. The collar is never 
complete below, always interrupted in the median line of the fore- 
neck by a narrower or broader dark longitudinal stripe between 
throat and breast ; iu one specimen ( 2 ^^-i Ishigaki, July; 5.11 .3.1)) 
this dark area is so broad as to occupy almost the whole width of 
the foreneck. Not infrequently the collar is partly interrnpted also 
on the back of the neck, owing to the pale tips being too short to 
completely cover the dark brown bases of the hairs ; in this case 
the " collar " is confined to the sides of the mantle, and sides of the 
neck and foreneck (compare Pi. fori)iosi(s). — Head varying, like 
back, from seal-brown to mars-brown, never uniform, but the 
amount of greyish and buffy admixture variable. 

The variations described above are independent of the sex of the 

Young female, scarcely half-grown (Ishigaki ; 5. 11. 3. 10): Darker 
and more uniform in colour than adults. Back, rump, breast, bellv, 
and flanks dark vandyck-brown, approaching seal-brown. Head 
blackish seal-brown. Mautle and sides of neck ochraeeous-butf. 

Measurements. On pp. 170. 171. 

Specimens e.vamined. The British Museum material, and the type 
of the species. 

Range. S. Liu-kiu Islands : Ishigaki, Iriomoto, Yonakumi, 

Type in the Leyden Museum. 

History in literature. — In one of the earliest known lists of 
Japanese mammals (Siebold, 1824) this species was recorded, under 
the name Pt. rviri/^Uis Geoff., from Nagasaki (Kiushiu) and Je<ld<t 
(Hondo). One year later (IB^.'i), on the atrcngth ol' a .single 
specimen, acquired by the Leyden Museum from Blomhoff, " ancicn 
resident ncerlandais au Japon," Temminck namcnl, described, and 
figured it as Pt. dasymaUus (the specific name in allusion to the 
dense woolly fur). Temminck copied Siebold's statement as to the 

162 J>TEROrUS IlASYMil.rXS. 

range of tlie sj.ecieB : Nagasaki and Jeddo, '• oil elle devaste les 
plantations d'arbres fruiticrs " ; in 1842 (Fauna Japonica) he 
wrote, however, that Pt. dastjmalhis occurs " seulement dans les 
parties meridionales de Tile Kiushiu, dans le district de Satsuma, et 
plus rarement a Jalamoshima.'' All records in literature of ditsy- 
mallns from Jeddo, Nagasaki, and Yakushima (see list of references 
above) are based exclusively on Temminck's statements ; in recent 
times the species has only been obtained in the S. Liu-kiu Islands ; 
one of its Japanese names, " Liukiu-komoli,'' i. c. Liu-kiu bat, 
would also seem to indicate that it is a native of this group of 
islands rather than of Japan proper. There can be little doubt, 
therefore, that Siebold was mistaken as to the habitat of da'symaJhis ; 
very likely he saw and obtained this species in Kiushiu and Hondo ; 
it is frequently brought alive to Japan from the Liu-kiu Islands 
and kept in captivity by the Japanese. 

The type in the Leyden Museum is an immature (not quite full- 
grown) mounted individual, skull extracted. The surface of the 
fur, above and beneath, has faded to whitish or creamy whitish ; 
base of hair less bleached. Skull and teeth as in British Museum 
specimens from the Liu-kiu Islands. 

Bj' (xraj' (1870) this species was placed in the genus Sjjectruni, 
between S. rubricoUe and S. anetianum. Though very different 
in skull and dentition, Pt. dnsi/malhis and anetianus are, in fact, 
externally so puzxlingly similar to each other as to be sometimes 
difficult to distinguish by external characters alone. 

(1. 5 imui. al. ; 'Sir E. Belcher Not reg. 

skull. L^'.]. 

b. c? ad. St. ; l^kllll. "Japan." Purchased ) 

(Verieaux). ( 
c-e. 1 J ad., 2 c? vg. Liu-kiu Is. H. Prver, Esq. 

ad. al. ■ [<-'•"]• 

/-!. 1 c? ad., 3 5 ad. Isliigaki, S. Liu- Purchased 5.11. .3.6-9. 

sks. ; skulls. kiu lf>. ; June- (A.Owston). 

July, 1904 (na- 
tive collector). 
/. 5 pull. sk. Ishigaki (native Purchased - 

collector). (A.Owston). 

k, I. c? ad., 2 ad. sks. : Iiiomoto, S. Liu- Purchased,12. 

skulls. kiu Is. : June (A.Owston). 

19 & 24, 1904 
(native collec- 
»H. $ad, sk. : sjaill. Yonakunii,S.Liu- Purchased 5.11.3 14. 

kiu Is.; June (A.Owston). 
23,1904 (native 
n. J; skull. Kobama, S. Liu- Purchased 

kiu Is.; July 20, (A.Owston). 
1904 (native 
collector). ' - 


lo. Pteropus formosus, P. L. Sdatcr. 
Puropus formosus, Dobson, Gat. Chir. E. M, p. 26. 

Pteropus formosus, P. Z. Sclater, P. Z. S. 1873 (Feb. 18), p. 193, 
pi. xxii. (animal) (Taku, Formosa) ; Gullioer, P. Z. S. 1875, 
p. 493* (size of red blood-corpuscles); Dohson, I. s. c. (1878: 
Formosa) ; Truuessaii, Rev. cS" Mag. Zovl. (3) vi. p. 204 (1879 : 
Formosa); P. L. iSclater, P. Z. S. 1891, p. 677 (Formosa); 
id., P. Z. S. 1892, p. 1 1 (Formosa) ; id., List An. Zool. Sac. 
Gard. 9 ed. p. 102 (1898)^ Trouessari^ Cat. Mamm. i. p. 78 
(1897 : Formosa). 

Pteropus (Spectruna) formosus, Matschie, Megachir. p. 27 (1899) ; 
Trouessart, Cat. Mainm., Suppl. p. 53 (1904 : Formosa). 

Diagnosis. — Similar to Pt. dasynudlns., but fur somewhat longer, 
colour almost uniform vandyck-browu above aud beneath, with 
creamy-white mantle and sides of neck. Forearm 130-137 mm. 
Hah. Formosiv. 

Skull awl teeth. — Xot differing appreciably from those of Pt. 

Palate-rithjes (two specimens, cotypes of species). — + 5^-3; 
sixth ridge in one specimen conspicuously, in the other verj- slightly 
interrupted in median line. Arrangement as in Pt. dasipnalhis. 

Far.— Longer than in J'l. dasymalJus : length of hair at middle 
of back about 29, middle of mantle 30, middle of belly 2S mm. 
Distribution and character of fur as in Pt. dasymaUits, 

Colour.— 2 ad. skin, Formosa, teeth slightly worn; B. M. Back and rump almost uniform vandyck-brown. Con- 
cealed bases of hairs vandyck-brown, tips paler, approximately 
mars-brown, or this colour with a slight touch of buffy. — Breast, 
belly, aud flanks vandyck-brown of a slightly darker tinge than 
back. — Collar represented by a large well-defined patch of creamy- 
white hair on either side of neck, and similarly coloured tips to I he 
hair of the mantle. Base of hair of mantle between vandyck-browu 
and seal-brown, tips of hair creamy white or silvery white, making 
a total impression of dark brown conspicuously varied and powdered 
with creamy-white. Hair of creamy-white patch on side of neck 
seal-brown only at extreme base, next to skin. Broad median portion 
of foreneck, between side patches, similar to breast. — Occiput, crown, 
and interocular space dark brown, between vandyck-brown and 
seal-brown, grizzled or pou dered with pale greyish ; sides of head 
more uniform dark brown; chin and throat almost seal-brown. 

One of the cotypes of the species, also a female (al. ; known to 
have been living almost seven years in continementj, is similar in 
colour to the specimen described above. In both of these females 
the mantle is mixed creamy-white and brownish, not defined by a 
straight line behind, but passing rather irregularly into the dark 
brownish colour of the back. In the second cotype, a male (al. ; 
almost seven years in confinement}, the whole of the mantle, sides 

* Miswritten Pt . J'oruto.-^aji ik<. t Misprinted Ft. mnrfiis. 


of neck, and sides of foreneck are unmixed crenmy white, forming 
a collar ver)- sharply defined from and contrasting with the hack, 
and interrupted only for a rather narrow space on the middle of 
the foreneck ; concealed base of hair of mantle, as usual, seal- 
brown. Similar variations in the colour of the mantle are seen in 
the closely related Ft. dasymallus (above p. 161) ; in both species 
they depend on the greater or less length of the pale tips to the 
hairs ; if long, the pale-coloured tips completely cover the dark base 
of the fur, making the mantle (in ft. /o?'?hosms) appear pure creamy- 
white ; if short, the dark base of the fur shows more or less 
through, making the colour of the mantle mixed dark brown and 
creamy-white, much less contrasting with that of the back. More 
material is required to decide whether in the present species these 
differences are merely individual (as is the case in Pt. dusymaJlns), 
or perhaps sexual. 

Measurements. On pp. 170, 171. 

Specimens examined. Three, in the collection of the British 

Range. Formosa (Taku). 

Coiypes in collection. 

History in literature. — A male and female of this species were 
presented to the Zoological Society's Gardens, Jan. 9. 1873, by the 
Kev. Mr. llitchie, of Taku, Formosa. The male was kept alive till 
October 4th, 1879, the female died presumably a few months later. 
A coloured plate of these two specimens was published, in 1873, by 
Sclater (Z. s. c), who named the species Pt. formosus, without de- 
scription; the specific name probably was jnoposed chiefly on account 
of its similarity to the name of the habitat of the species. The 
plate does not give a correct idea of the colour of the fur. The 
earliest description is that published by Dobson, in 1878. A third 
specimen, also from Formosa, was presented to the Society's Gardens, 
Dec. 1, 1891, by Mr. Thos. Perkins. The skin of a female speci- 
men, according to a note on the label, " brought alive, said to have 
come from Formosa," was ))resented to the British Museum, in 
1892, by Mr. J. de La Touche. So far (December, 1908), these 
seem to be the only examples on record. 

a. S ad. al. ; skull. Taku, Formosa Zool. Soc. Gardens. 80.0. 25. 7. 

h. 5 ad. al. ; skull. Taku, Formosa Zool. Soc. Gardens. 
_ [Ritchie). 

(a, b : Cott/pes of species.) 
c. 5 ad. sk.; skull. Formosa. J. de La Touche [P.J. 

16. Pteropus subniger, Kerr. 

Pteropus rubricolUs, Dobson, Cat. Chir. B. M. p. 24. 

La Roussette a col rouge,- Brisson, Rm/h. Anim. p. 217 (1756: 

Reunion) ; id., op. cit. 2 ed. p. 154 (1762: Reunion). 
La RougettP, Btiffon, Hi't. Aaf. x. pp, 5.5, 7fl, 82, pi. xvii. (animal) 


(1763 : Reunion) ; id., op. cit. Suppl. iii. p. 256 (1776 : lleunion ; 
habits) ; Fouchet d'Obsunville, Essais philosophiques, p. 77 (17(:'3; 
Keuniou) ; G. Cuvier, Tahl. Elem. p. 104 (1798) ; E. Geoffroij, 
Ann. Mas. iVHist. Nat. vii. p. 230 (1806 : Mauritius ; habits). 
Ternate Bat : The Hom^ettc, Pennant, Hist. Quadr. ii. p. 549 (1781) : 

id., op. cit. 3 ed. ii. p. 305 (1793). 
Eoussetto ; une autre espeee, Milbert, Voy. pittor. Ik-de-France 

p. 245 (1812 : Mauritius). 
Vespertilio vauipyrus (iiec L.), P. L. S. Miiller, Vollst. Naf.ursyst. i. 
p. 153 (pt.) (1773: Reunion). Var. B, Schreber, Sdiic/. i. 
pp. 153, 155 (1774); Boddaert, Eleiich. Anim. i. p. 68 (pt.) 
(1785). Var. ji, Gmelin, Linn. 8yd. Nat. i. p. 45 (1787). Var. 3, 
Turton, Linn. Syst. Nat. i. p. 24 (1802). 
Pteropus vampyrus, var. fi, Er.vlebcn, Syst. Regn. Aiiim. p. 133 
(1777). Var. /3, Bonndorff, Zool. Beytr. i. p. 62 (1792). Var. b, 
(Die Euugette), Bechstein, Pennunfs Vicrfuss. T/iicre, ii. pp. 617 
733 (1800). 
Vespertilio vampyrus subniger, Ken; Anim. Kinyd. i. pt. i. pp xvii 

91, no. 107 (1792). 
Spectrum rubidum, Daudin, in Biiffon, Hist. Nat., Didof, ed Quadr 

xiv. p. 188 (1802*). 
Pteropus ruber, E. Geoffrey, Cat. Mamm. Mus. Nation. d'Hist. Nat. 

p. 48 (1803 : Madag'iiscar, errore). 
Pteropus fuscus (wee E. Geof.) Desmarest, N. Diet. d'JIist. Nat. 

xix. p. 544 (1803 : Reunion). 
Vesperiilio caninus {nee Blumenb.), var. a, Goldfuss, Very/. Naiur- 

beschr. Sduy. p. 98 (1809). 
Pteropus rubricollis, E. Geufroy, Ann. Mus. d'JIist. Nat. xv. p. 93 
(1810: Reunion); Oken, Le/irb. Naturg. iii. Abth. ii. p. !t35 
(1816: Reunion ; Mauritius) ; G. Cucier, Rkjne Anim. i. p. 124 
(1817: Mauritius; Reunion); Desmarest, 'Mamm. i. p. 110, 
no. 140 (1820: Mascarenes) ; Schinz, Thierr. i. p. 154 (1821 : 
Mauritius; Reunion) ; Temminek, Mon. Ma^nm. i. p. 183 (1825: 
Reunion; Madagascar); Lesson, Man. Mamm. p. 109, no. 280 
(1827 : Reunion) ; Gray, Griffith's Anim. Kinyd. y. p. 55, no. 156t 
(1827 : Mauritius) ; Desmarest, Diet. 8ci. Nat. xivi. p. 362 
(1827: Mauritius; Reunion); Is. Genffroy, Diet. Class. d'Hist. 
Nat. xiv. p. 700 (1828: Reunion) ; J. B. Fischer, Syn. Manm. 
p. 84, no. 8 (1829 : Reunion) ; Temminek, Mon. Mamm. ii. p. 75 
(1837); Waterhouse, Cat. Mamm. Mus. Zool. Soe. Land. f. 13, 
no. 102 (1838: Mauritius) ; Gray, May. Zool. ^- Bof. ii. p. 503 
(1838: Cape of Good Hope; Reunion; Madagascar) ; Oken, 
Ally. Natury. vii. Abth. ii. p. 988 (1838: Mauritius; Reunion; 
Madagascar) ; Wayner, Schreber's Sduy., Suppl. i. p. 351 (1839: 
Reunion ; iladagascar) ; Lesson, N. Tabl. Regne An., Mamm. 
p. 13, no. 178 (1842 ; Reunion ; ]\Iadagascar ; CafTraria) ; 
Schinz, Sysf. Verz. Sduy. i. p. 125 (1844: Reunion; Mada- 
gascar; Caffrariii); Wayner, Schreber's Sdug., Si/ppl. v. p. 602 
(1853-55 : Reunion ; Madagascar) ; Gervais, Hist. Nat. Mamm. 
i. p. 188 (1854); Giebel, Sduy. p. 997 (1855: Reunion; .Mada- 
gascar) ; Peters, MB. Akcid. Berlin, 1867, p. 323 (Reunion) ; 
Fitzingcr, SB. Akad. JVien, Ix. Abth. i. p. 453 (1870: Reunion : 

* I'itlepns^e dated 1799. On the true Tear of publication see W Eich- 
n.ond, The Auk, xvi. p. 529 (1899). 
t Misspelt Pi. ri«6iC(j//is. 


Mauritius) : Miirchi, Atti Soc. Ital. Sci. Nat. x\. p. 516 (1872-73: 
structure of hairs) ; Uohsuii, Cat. Chir. B. M. p. 24 (1878: 
Mauritius) ; Troue.ssarf, Rev. ^ Maff. Zool. (3) vi. p. 204 (1879 : 
Jrienuion ; Maiuitius) ; Rohin, Ann. Sci. Nat. (6) Zool. xii. Art. 2, 
p. 4 et seq. pi. ii. tig'. 1, pi. iii. Ho-. 16 (1831 : anatomy) ; Troues- 
sart, Cat. Mavtm. i. p. 77 (1897: Keuuion ; Mauritius) ; Mntschie, 
Meyachir. pi. viii. tiifs. 1, 1 «. \a [bis], 1 6, 1 e (bead, teeth) 
(1899 : >rauritius) ; Elliot, Cat. Mamm. Field Col. Mus. p. 491 
(1907 : Mauritius) ; Miller, Fain. ^ Gen. Bats, p. 58 (1907). 

Spectrum rubricolle, Gray, Cat. Monk. Src p. 101 (1870: Cape 
of Good Hope; Mauritius). 

Pteropus (Sericonycteris) rubricollis, Matschic, Megachir. p. 30 
(1899: Reunion; Mauritius): Trone»sart, Cat. Mamm., Suppl. 
p. 54(1904: Reunion; Mauritius). 

Pteropus torquatus, G. Fischer, Zoof/nosia, iii. p. 553 (1814). 

Pteropus collaris, Illif/er, Ahh. Akad. Berlin, 1804-11, pp. 78, 84 
(1815: E. Africau Islands). 

Diar/nosis. — Teeth much smaller, especially narrower, than usual. 
Ears small, concealed in the fur. Tibia densely clothed above. 
Fnr of mantle, back, and rump long, scmicrect. Orange bufTy 
collar contrasting with general dark brownish colour of fur above 
and below. Size very small : forearm 95-99 mm. Hab. Mas- 

SJadl. — SfDall, delicately built, with short rostrum. Brain-case 
high, domed, profile-line of skull from top of brain-case to nasals 
therefore steeper than in Pt. hypomdanus. Deflection of brain- 
case greater than usual, the alveolar line if projected backward 
passing through middle of vertical portion of supraoccipital. The 
shortness of the rostrum chiefly due to the reduction of the teeth 
and the comparatively larger ute of the orbits ; front of orbit 
vertically above back of p'; orbital diameter only slightly less than 
lachrymal width of rostrum. Sagittal crest undeveloped ; the 
temporal crests in fully adults closely approximated to each other, 
but apparently never fused in the median line to form a sagittal 
crest. Postorbital processes long, slender, reaching about halfway 
(or a little more) between frontal and zygoma ; corresponding 
processes on zygoma very small or undeveloped. Coronoid height 
of mandinh^ much less than length of lower tooth-row, c-ffig. 

Tietli. — Chief character : cheek-teeth small and unusually 
narrow ; width of p* between one-fourth and one-fifth (in most 
other species of the genus about one-third) the palatal width 
between p^-p''. 

Cingulum of upper canines strong, shelf-like, p' minute, terete, 
spiculiform, decidtioiis : in six almost full-grown individuals it is 
present in five, lost (and alveolus closed) in one; in two adults 
■with unworn teeth present in one, lost in the other. Posterior 
ledo-es of p^ and p* feebly developed, without postero-external cusp, 
or with only an indication of such in p'. m^ not quite twice as 
long as broad, m- nearly equal in size to p,, larger than m,. — 
i in cross section once and a half or scarcely twice the size of i^. 
tiiigulum of lower canines M'ell developed, but narrower than in 


upper canines, p, rather large, mucli larger than m^, about twice 
(or a little moi'e) the bulk of i„. Posterior ledges of pj and p^ well 
developed, raised postero-externall)' into a minute cusp; a similar 
cusp-like elevation sometimes faintly indicated iu m,. m., small, 
subequal in bulk to i„. 

The remarkable reduction in the siiie of the teeth accounts for 
the non-development of a sagittal crest in this species. 

Ears. — Very small, completelj' bidden in the fur ; triangular in 
shape, outer and inner margins almost evenly converging I'rom 
base to tip. Densely clothed with long hairs from base to tip, in 
front and behind. 

Interfemoral. — Extremely short or scarcely developed in centre. 

Far, — Long, very dense, silky, somewhat crinkled ; semierect 
over the whole of the upperside of the body, directed backward but 
not closely adpressed on underparts. Length of hairs at middle of 
back about 25, middle of mantle 2fJ, middle of belly '22 mm. 

Above, proximal half or third of forearm clothed with long, 
dense, adpressed hairs. Fur of back extending laterally on the 
membrane almost to a line between middle of forearm and knee. 
Tibia long-haired to ankle ; upperside of metataKUs and phalanges 
to base of claws with long, thinly spread hairs. Below, proximal 
half of tibia covered with long, dense fur. 

Colour. — Ad. skin, Mauritius, B. M. Eack and rump 
blackish seal-bi'own ; a slight sprinkling with silvery greyish-wliite 
hairs detectable only on close examination. In the front of the 
back, near the mantle, the individual hairs are seal-brown at 
extreme base, then broadly buff, tip seal-brown ; further backward 
the middle pale-coloured portion of the hair is gradually becoming 
narrower and more greyish, until at middle of back ifc is represented 
only by an indistinct greyish powdering of the middle ] ortion of 
the hair, Avhich apart from this might be described as uniform 
seal-brown ; on the rump the pale-coloured ring is again becoming 
more pronounced and broader, passing from approximately mouse- 
grey to smoke-grey near the interfemoral ; all pale-coloured parts 
of the hairs of the back and rump only detectable on separating the 
fur. Hair on upperside of tibia pale bufiy or cream-bufty. — 
Breast, belly, and tianks seal-brown, sparsely and inconspicuously 
sprinkled with some long buffy hairs. Anal region next to intei- 
femoral and fur on underside of tibia bufl'y or cream-buffv. — 
Alantle rich orange-buff, all the hairs seal-brown at extreme base. 
On the sides of the neck and foreneck the colour gradually darkens 
to tawny ; extreme bases of hairs evcnwhere seal-brown. — Inter- 
ocular space, crown, occiput, and sides of bead grizzled dark 
brownish and greyish buiTy ; individual hairs seal-browu at extreme 
base, then broadly cream-buffy or (jreyish creain-buffy, seal-broAvn 
at extreme tip; the general grizzled aspect of the colour of the 
head being duo to the fact that the seal-brown tips are too short to 
completely conceal the paler middle portion of the hairs. Chin and 
throat uniform seal-brown. 


IiuliviJual variation in colour :- — Eack and ruiup in some speci- 
mens leas blackish (more typical) seal-brown, in others more 
'• powdered " with greyish, chierly owing to abrasion of the hair- 
tips, by which the paler middle portion of the hairs gains more 
influence on the general aspect of the colour. Tibia above and 
beneath often similar in colour to back, without, or with only an 
indistinct, tinge of bufty. — Underparts often more conspicuously 
sprinkled with long pale-coloured hairs.— Colour of mantle varying 
from ochraceous-bufl' to orange ochraceous and tawny ochraceous. 

Measurements. On pp. 170, 171. 

Specimens examined. Ihe material in the Paris, Berlin, and 
iJritish Museums. 

Range. The Mascarenes : Keunion, Mauritius. 

Type probably not in existence. 

Hahits. — Ft. suhniijer is said to be strictly nocturnal in habits. 
It hides in holes in trees or in caves, more than four hundred 
being sometimes found together. It is popularly believed in 
lleunion that however large is the number of Kougettes found in a 
cave, there is always only one male ; this probably means that; 
apart from the rutting-season, the sexes of this Fruit-bat, like 
many other bats, occupy separate resting-places. (De la Nux, iu 
liuffon's f^ijppl. iii. p. 250, 1776; lloch, quoted by GeotFroy, Ann, 
Mus. d'Hist. Nat. vii. p. 230, 1800.) 

Earliest histori/ in literature. — Owing to its exclusively nocturnal 
habits and its hiding in hollow trees and eaves, this species, though 
once common in Eeunion and Mauritius, was not quite so well known 
to the French zoologists in the latter half of the eighteenth century 
as Ft. niger. It is perhaps Daubenton's "lioussette" (Mem. Acad. 
Sci. 175y, p. 385). Brisson's excellent diagnosis (1750 and 1762) 
of the 'lioussette a col rouge ' (fuscus, auriculis brevibus acutiusculis. 
collo superiore rubro) was taken from a lleunion specimen in the 
Cabinet lleaumur. Butfon's description and figure of the Kougette 
(1763) were based on a dried specimen in the Royal Cabinet, sent 
from lleunion by de la Nux, " ancien Conseiller au Conseil royal de 
cette lie," his later account of its habits (1776) on information from 
the same observer. Pennant (1781) put the llougette of French 
writers down as a variety of Seba's " Ternate Bat." 

Linne, in his 10th and 12th editions (1758 and 1766), has no 
reference to the llougette. The early post-Linnean compilers 
(P. L. a. Miiller, Erxleben, Boddaert, Gmelin, Donndorfi') record 
tlie species as a variety of the Linnean Vcspertilio (or Fterojms) 
vampyrus. The same view was taken by ISchreber (177-1), who 
probably knew the species only from literature. 

Vespertilio varnpyrvs subniger, Kerr; 17^2. — Based on Brisson's 
Pteropxis collo ruhro. Kerr's brief description is taken partly from 
Brisson, partly from BufFon, his notes on its habits from de la Nux's 
letter to Buflbn. Inasmuch as Brisson's Ft. collo rubro and Buffbn's 
llougette were based on a specimen from lleunion, this island must 
be fixed as the type locality of Ft. s^ihmger. 


Spectrum rubidum, Daudiii ; 1802. — Name proposed by Daudin 
(in his revised edition of Lacepede's Tabl. Meth., published in the 
IJidot Biitfou) for Biiffon"s llougette ; type locality therefore 

Pteropus ruber, ¥..Oeo&voj ; 1803. — Geoffroy's brief description 
of Ft. ruber leaves no room for doubt as to the identification. The 
references given by GeofTroy are Pi. fuscus Brisson (i. e. Pt. fuscus, 
auriculis brevibus acutiusculis, coUo superiore rubro), the Roussette 
of Daubenton, and the Bougette of Buffon, all of which (or at all 
events the first and third of which) are Ft. subniger. The single 
specimen oi Ft. ruber catalogued by Geoffroy (his no. 91) was stated 
to have been " envoyc do Madagascar par le citoyen Mace." This 
error as to the locality was repeated, in 1825, by Temminck (who 
is known to have possessed a copy of the rare " Cat. Mamm. Mus. 
Nation. d'Hist. Nat.,"' which was never offered for sale but 
I)rivately distributed to a good number of zoologists in France 
and abroad ; Temminck's copy is still in the Leyden Museum); all 
later writers' records of the species from Madagascar rest solely on 
this mistake. — The Paris Museum does not now possess any specimen 
Avliich can be identified with the type of Pt. ruber. 

Fteropus fuscus, Desmarcst ; 1803. — Type locality, Reunion. 
Only references given by Desmarest : Bufibn's llougette and 
Brisson's Fteropus fuscus, Roussette a cou rouge, both of which are 
Ft. subnujer. The name ^^ fuscus'" was evidently bori-owed from 
Brisson's diagnosis of Fteropus collo rubro (" Pt. fuscus, auriculis 
brevibus " &c.). — Desmarest's Ft. fuscus should not be confused 
with Pt. fuscus, E. Geoff., 1803, which is Ft. nigcr. 

Pt. rubricollis, E. Geoffroy ; 1810. — Type locality, Reunion. 
Based on one specimen, in the Paris Museum, formerly in the 
Cabinet Reaumur, sent by de ]a Nux. This specimen was no doubt 
the verj' typo of Brisson's Fteropus collo rubro (see above), probably 
also the example described and figured by Bufibn as the " Rougette." 
It is no longer in existence. 

Pt. torqvatus, G. Fischer; 1814. — A renaming and redescription 
of Brisson's Ft. collo rubro. Type locality therefore Reunion. 

Fteropus collaris, IWigcr ; 1815. — Based, without description, on 
Buffon's Rougette. Typo locality therefore Reunion. — In 1828 
(Verz. Doubl. Mus. Berlin, p. 3, no. 47), Lichtcnstein wrongly 
identified llliger's Pt. collaris with the S. African Fruit -bat 
later on (1829) described by A. Smith as Fteropus leachi (i. e. 
liousettus leachi of this Catalogue), giving as habitat of Pt. collaris 
" Terra CafTrorum." Some statements in literature of the occur- 
rence of Ft. rubricollis in S. Africa (Lesson, 1842 ; Schinz, 1844 ; 
Gray, 1870; 1. s. c.) are undoubtedly based on this error of 

Femarks. — Tlie following three characters taken together aro 
sufficient to distinguish Pt. subniger from any other species of the 
genus : Cheek-teeth much reduced in size ; tibia densely haired 
above ; fur dark brownish above and beneath, with orange ochraceous 



collar. The first character distinguishes it from all species of 
Fteropus except molossinus, persoaatus, scapulatus, and woodfordi : 
the second excludes molossinus, scapulatus, and woodfordi ; the 
third, personatus. 

a. Inini. sk. ; skull. 

b. (5 imm. st. 

c. Imm. sk. ; skull. 

d. (S yg. ad. sk. ; 


e. Imm. sk. ; skull. 
f.g. 2 imm. sks. ; 

h, i. 1 ad., 1 2 iuiui. 
sks.; skulls. 

Leydeu Museum. 
J. Gould, Esq. [P.], 

Purchased (Parzudaki). 
Tomes Coll. (Zool. Soc). 

Mauritius. H. Whitely, Esq. [P.]. 

Not reg. 
Not reg. 
37.4.28 31., 
257 bis., .3. 

External measurements of Pteropus dasymallus, formosus, 
and subniarer. 


Polles, total length, c. u 

2nd digit, metacarpal 

„ 1st phalanx 

„ 2nd-3rd phalanx, c. u 
3rd digit, metacarpal 

,, 1st phalanx 

,, 2nd phalanx 

4th digit, metacarpal 

,, 1st phalanx 

,, 2nd phalanx 

5th digit, metacarpal 

,, 1st phalanx 

,, 2nd phalanx 

Ear, length from orifice 

„ gi'eatest vidtb, flattened .. 
Front of eye to tip of muzzle . . 

Lower leg 

Foot, 0. u 


Pt. dasymallus. 
12 ad. 

Mil*. Max. 














45 5 








Pt. formosus. 

3 ad. 
(Inch cotypes.) 

MiN. Max. 

58 -o 

















P(. suhniger. 
3 ad. 

MiN. Max. 
































Meusartinents of skulls and tuth of Pteropus dasymallus, 
formosus, and subniger. 

Skull, total length to gimtbion 

,, palation to incisive foramina 

„ front of orbit to tip of nasals 
„ width of brain-case at zygomata.. 

„ zygomatic width 

„ width acrofs ni', extei-nally 

„ lachrymal width 

„ width across canines, externally.. 

„ postorbital constriction 

„ interorbital constriction 

„ width of me.'opterygoid fossa 

,, between p'^-p^, internally 

,, orbital diameter 

Mandible, length 

„ coronoid height 

Upper iceth, c-m'- 

Lower teeth, c-m, 

Upper incisors, combined width 

p*, Isngth 

,, width 

P'', length 

„ width 

m", length 

„ width 

Ill', length 

„ width 

Pi, length 

„ width 

p,, length 

I „ width 

i P«. length 

, „ width , 

m,, length 

„ width 

irij, length 

„ width 

m„ length 

I I, width 

Ft. dasi/mallus. 

Skulls : 9 ad. 

Teeth : 9 ad., 

1 imm. 

(Incl. type.) 

Mi.-i. Mas. 

Ft. foi-mosus. Ft. subniger. 

Skulls: Sad. ] Skulls: 2 ad. 

Teeth : 3 ad. ' Teeth : 2 ad., 
(Incl. cotypes.) 1 6 imm. 

































MiN. Max. 













AIiN. Max. 

























































B. TuE Pteropis mariaxxv.s group. 

Species. — Eight, Ft. iJeleiuensis, yapensis, ualaaus, mariannus, 
loochoensis, vanikorensis, tongamts, and r/cddiei. 

Range. — Polynesia (so far as inhabited by the geuiis), extending 
north-west to South Liu-kin Islands. 

General characters.- — Skull and dentition unmodified Pteropine ; 
posterior basal ledges of premolars slightly more developed than in 
Ft. Inipomelanus. General shape of ears and distribution of fur as 
in Pi. h)/pomelanus ; ears moderate, broad, tip rounded off; fur 
short, adpressed on back ; tibia naked above. Colour pattern 
remarkably constant : in nearly all species, mantle light yellowish 
strongly contrasting with blackish or dark brownish back, muzzle, 
and underparts ; dark colour often sprinkled with greyish or 
whitish, especially on underside. Sexual differentiation inconsi- 
derable : males without glandular neck-tufts, but generally with 
rather heavier canines and fur of mantle more rigid, oily, and 
uniform buffy (in females softer, more spreading, and with concealed 
brownish bases to the hairs). Size generally moderate or rather 
small (forearm 113-154 mm.). 

Differentiation of species. — The eight species are separable into 
two sections : a series of relatively smaller-eyed North Pacific forms 
(/*<. pelewensis, yapensis, ualanus, mariannus, and loochoensis), dis- 
tributed over the Pelew, Caroline (two species), Marianne, and South 
Liu-kiu islands ; and a series of relatively larger-eyed South Pacific 
forms {Ft. vanil-orensis, tonf/anus, and geddiei), ranging over the Santa 
Cruz, New Hebrides, New Caledonia, Fiji, Tonga, and Samoa islands. 
Ft. peleiuensis is a peculiarly small and small-toothed species, in the 
colour of the fur rather closely approaching Ft. admiralitatum. The 
four other North Pacific species are closely interrelated ; from 
Ft. peleivensis they differ chiefly in their larger size, from each other 
in the size of the teeth and length of the fur ; the teeth are smallest 
in Ft. ualanus (East Carolines), heavier in Ft. yapensis (West Caro- 
lines) and Ft loochoensis, rather heavier still in Ft. mariannus ; the 
fur is longer than usual in the extreme north-western species, 
Ft. loochoensis. — Also the three species of the South Pacific section 
are closely related. Ft. vanikorensis (known from Vanikoro only) 
is chiefly characterized by its shorter. Ft. tonganus (P'iji, Tonga, 
and Samoa islands) by its longer wings ; Ft. geddiei (Now Hebrides 
and New Caledonia) is the largest form of the group. 

Affinities of group. — The Ft. mariannus replaces the Ft. hypo- 
melanus group in Polynesia ; tliere can be little doubt that it is a 
Polynesian modification of the hypomelanus type. Taken as a whole 
it differs from this latter only in characters of trivial importance : 
the slightly heavier posterior basal ledges of the premolars and the 
style of colour. The former character indicates scarcely more than 
a slight difference of degree ; the style of colour is closely approached 
by a species of the hypomelanus group, Ft. admiralitatum ; and both 
characters are less conspicuous or, as it were, less definitely fixed 
in the species which geographically are neighbours to the area of 


the hjipomelaims group. It is, in fact, open to question wLetlior 
Pt. admiral'datum ought to be considered a peripheral species of the 
liypomelatms type with strong leanings to the Polynesian group, or 
rather incorporated in this latter. — The species of the Austro- 
Malayaii and Australian Pt. conspicillafvs group (P. clirysauchen, 
conspiciUatus, ncularis) are almost precisely similar in colour to the 
typical members of the marlamius group(compare e.g. Pt.chnjiauchen 
^vith Pt. tongainis) : any close relationship between these two sections 
of the genus is, in view of the difference in dentition, altogether 

17. Pteropus pelewensis. A'. And. 
Pteropus heraxulreni (pt.), Dobson, Cat. Chir. B, M. p. 63. 

Pteropus keraudreni {nee Pt. kevaudren, Q. 4' G.), var. a (pt.), 
Dobson, Cat. Chir. B. M. p. (>5 (1878: Pelewls.). Yar. aa (pt.), 
Trouessart, Rev. ^ Mat/. Zuol. (3) vi. p. 203 (ls79 : Pelew Is.). 

Pteropus insulavis (pt., nee Hombr. Sf Jacq.), Trouessart, Cat. Mamm. 
i. p. 83 (1897: Pelew Is.). 

? Pteropus (Spectrum) mariannus (pt., nee Desm.), MatKchie, Mega- 
chir. p. 27 (excl. figs.) (1899: Pelew Is.); Trouessart, Cat. 
Maynm., Siippl. p. 53 (1904 : Pelew Is.). 

Pteropus pelewensis, E. Andersen, Ann. ^- Mao. N. II. (8) ii p 364 
(1 Oct. 1908: Pelewls.). 

Diarjnosis. — Allied to Pt. admiralitatum (supra p. 144), but with 
much smaller orbits, narrower rostrum, weaker dentition, and shorter 
fur. Colour of fur approaching that of Pt. admiralitatum. Forearm 
shorter; 113-5 mm. Hab. Pelewls. 

Skidl and teeth. — General size of skull nearly as in Pt. admirali- 
taiwn, but rostrum considerably narrower and orbits smaller: 
maxillary width externally across m'-m' 14-8 mm., against 1(5-17 

in Pt. admiralitatuni ; orbital diameter 11 mm., against 12-12-5. 

Structure of teeth as in the allied species, but dentition on the 
whole slightly weaker, p^ and m^ markedly smaller ; see measure- 
ments, p. 176. 

Fur. — Length of fur as in Pt. hijpornelanus, much shorter than in 
Pt. admiraUtatum : on back 9-11, mantle 1:2-14, belly 13-14 mm. 
Distribution of fur as in the allied species. 

CWoH?-.— Type, cJ ad. skin, teeth much worn. — Back and rump 
dark brown approaching seal-brown, conspicuously washed with 
vandyck-brown on sides of back, humeri, and thighs, and thinly 
sprinkled with silvery greyish-white hairs. — Breast, belly, and 
flanks a dark shade of Prout's brown, everywhere thickly mixed 
with shiny silvery white or buify-white hairs. — Mantle and sides 
of neck light yellowish buff, individual hairs uniform from base to 
tip or, in shoulder region, with extreme base blackish. Foreneck 
similar to breast. — Occiput similar to mantle, but somewhat clouded 
with brownish. Crown and sides of head mixed buff'v and brownish 
sprinkled with some highly glossy silvery grey hair?. Chin and 
throat blackish thinly sprinkled with silvery grev. 


A young female (not full-grown, skin, differs conspi- 
cuously from the type in colour. Back and rump blackish thickly 
mixed with shiny silvery grey hairs or tips to the haii-s. — Breast, 
belly, and flanks blackish seal-brown heavily sprinkled with shiny 
silvery grey. — Mantle pale buffy cinnamon, shading into deep mars- 
brown on sides of neck, and into dark chocolate on foreneck ; base 
of hairs everywhere blackish. — Crown darker than mantle and 
sprinkled with glossy silvery grey ; colour gradually darlicniug on 
sides of head into the blackish seal-brown of throat. 

It appears probable (judging from allied species) that the differ- 
ence in the colour of the base of the fur of the mantle (yellowish 
buff like tips of hairs in the above-described male, blackish in female) 
is not dependent on age, but sexual. The hairs of the mantle 
are rather rigid in the single male examined, soft and spreading in 
the single female. 

Measurements. On pp. 175, 176. 

Specimens examined. Two, as registered below. 

Range. Pelew Islands. 

Type in collection. 

a. cJ ad. skin ; skull. Pelew Is. ; 1870 (Crt/'C. Godeffroy Miiseiim. 

Heinsohn). (.Type of species.) 

h. 5 jun. skin ; skull. Pelew Is. ; 1870 (C'ajo;'. Gocleffroj Museum. 


18. Pteropus yapensis, K. And. 
Pteropus keraudreni (pt.), Dobson, Cat. Chir. B. M. p. 63. 

Pteropus keraudreni (nee Pt. keraudren, Q. S,- G.), Tetens *.§• Kubanj, 

Journ. Mus. Godeffroy, i. pt. ii. p. 50 (122) (1873 : Yap). 
Pteropus keraudreni, var. a (pt.), Dohson, Cut. Chir. B. M. p. 65 

(1878 : Yap ; Macltenzie). Var. aa (pt.), Trotiessart, Rev. ^- Mag. 

Zool. (3) vi. p. 203 (1879 : Mackenzie). 
Pteropus iusularis {}^t.,ncc Homhr. ^-Jacq.), Trouessart, Cat. Mamm. 

i. p. 83(1897: Yap; Mackenzie). 
Pteropus (Spectrum) mariannus (pt., nee Desm.), Matschie, Meya- 

chir. p. 27 (excl. figs.) (1899: Yap; Mackenzie); Trotiessart, 

Cat. Mamm., Suppl. p. 53 (1904 : Yap ; Mackenzie). 
Pteropus yapensis. A'. Andersen, Ann. 4- May. N. H. (8) ii. p. 365 

(1 Oct. 1908 : Yap ; Mackenzie). 

Diagnosis. -i'>\ixi.\\d,v to Pt. admiraliioium, but somewhat larger, 
with broader skull, smaller orbits, more strongly developed posterior 
ledo-es of cheek-teeth, longer ears, and shorter fur. Blackish above 
and beneath, sprinkled with whitish ; mantle and sides of neck 
strongly contrasting, yellowish buff ; foreneck washed with russet. 
Forearm about 130 mm. Hah. W. Carolines. 

Skull and teeth. — Size of skull as in Pt. admiralitatum, if not 
slightly larger, but temporal fossa much broader, zygomatic arches 
therefore much more flaring posteriorly, sagittal crest stronger, 
frontal region between orbits broader, orbits smaller; zygomatic 
width about 3(5 ram., against 32-33 in Pt. admiralitatum ; orbital 


iM-cr),.,. ^, -1 ; Js'HiJ^t l.i;-l^o; coronoid process markedlv 

.e'rlva^inp/ ;;'"'"''', '"^"i""' *" c-m^-Size of teeth 
i.earlj as m Pt. admrrahtatura, though ,/ and p-' distinctly heavier 
(see measurement.s, p. 176), but posterior ledges of p3 p^p T 

postcro-exUnualh ; cmgulum of canines somewhat broader. 

A«; ..-Larger than m Pt. admlralUatum, very nearly as in Pt 
h,jpomelnnus ; shape as in these species ^ 

Interfemoral.-Hhon or undeveloped in centre 
back" 9I1 l^Jl'^ closely adpressed on back. Approximate length, 

S;i u '.. -' ^"^ ^^""^'^ seal-brown, darkest in mid le line 

£s?-an7tt"lt'T'"'^'^ paler shade of brown" n^diof 
oreasr and bell}, everywhere conspicuously sprinkled with elossv 

Sldef;:Nr:-r,';^^:-"^''^ hairl-Maitli and Midi f'nel 
ottr di tnIS wl'^'r T' P^'''°^^ aPP^-oachiug buff-yellow, t 
others distinctly washed with orange-buff, strongly contrasting ^ith 

Ea:ternal measurements o/Pteropus pelewensis and yapensis. 


Pollex, total length, c. u. ....... 

>> metacarpal 

j ,. 1st phalanx | 

I 2n(l digit, metacarpal ....'', 

I .. 1st phalanx 

,, 2nd-3rcl phalanx, c. u. 

I ord digit, metacarpal 

,. 1st ptialanx 

I ,. 2nd phalanx 

! 4tb digit, metacarpal 

' II 1st phalanx 

[ ,. 2nd phalanx 

! 5th digit, metacarpal 

I, 1st phalanx 

t. 2nd phalanx 

Ears, length from orific-e 


Lower leg 

Foot, c. 11 ] I 



Pt, peleweiisis. 

Pt. i/apeiim. 

d ad. 

6 ad. 







; 130 

1 58 


i 14 


' 29-5 


' 65 










98 5 









41 ' 








* Ksliniaie iskin). 



Measurements of skulls and teeth of Pteropns pelewensis 
and j'apensis. 

Skull, palation to incisive foramina ... 
,, front of orbit to tip of nasals . . . 
„ width of brain-case at zygomata. 

„ zygomatic width 

,, width across m\ externally 

„ lachrymal width 

„ width across canines, externally. 

„ postorbital constriction 

,, interorbital constriction 

,, width of mesopterygoid fossa ... 

,, between p^p*, internally 

,, between cingula of canines 

,, orbital diameter 

Mandible, length 

„ coronoid height 

Upper teeth, c-m^ 

Lower teeth, c-ni, 

Upper incisors, combined width 

p^ length j 

,, width 

p^, length 

,, width 

ml, length 

,, width 

m*, length 

,, width 

Pi, length 

,, width 

Pj, length 

,, width 

p,, length 

,, width 

mj, length 

„ width 

m^, length 

., width 

m3, length 

„ width 

Pi. pelewensis. 
Type * and 

Pt. yapensis. 
Type and 

S ad. 













f? jim. 





















4 1 






S ad. 

































2 jun. 




* Teeth much worn. 


and sharply iiiaikocl off from blackish back ; on the foreueck tho 
yellowish colour is considerably clouded with pale russet. liase of 
fur of mantle yellowish buff like tips of hairs, except posteriorly 
near shoulders where the hairs have sliort blackish bases. — Occiput, 
as far forward as a lino between ears, similar to mantle, but washed 
with russet at base of cars. Crown and interocular space mixed 
buffy and dark brown with blackish bases to the liairs and thinly 
sprinkled with silvery whitish. Colour gradually darkening ou 
sides of head, and passing on throat into blackish thinly sprinkled 
with silvery whitish. 

An immature female (skin, Mackenzie I., 7- is on tho 
whole similar in colour to the type, but with the geuei-al tinge of 
the hack and rump more approaching seal-brown than hlackish ; 
mantle and sides of neck unusually pale, cream-buff with long seal- 
brown bases to the hairs ; foreneck much clouded with seal-brown. 

tSe.vual differentiation. — Probably as in Pt. tonganus. The only 
male examined of Pt. yapensi'i has the hairs of the mantle somewhat 
rigid and uniform yellowish buff from base to tip (except poste- 
riox'ly, near line of demarcation between bright mantle and blackisli 
back). Tho only female is immature ; fur of mantle soft and 
spreading, everywhere with long dark brown bases to the hairs. 

Measurements. On pp. 175, 176. 

Specimens examined. Two, as registered below. 

Jiange. Western Carolines : Yap, Mackenzie. 

Tjipe in collection. 

fl. (fad. skin; Yap, W. Carolines (Cf^j/. Godoi'tVoy Muspum. 

skull. JWers). (Ti/pc of species.) 

i. 2jun. skin; Mackenzie, W. Carolines Grodctfroy Museum. 

skull. {Capt. Peterti). 

ID. Pteropus ualanus, Pet. 

Pteropua keraiidren (pt., nee Q. ^- G.), Lesson, Man. Ma mm. p. lOP, 

no. 281 (1827 : E. Carolines) ; id., N. Tabl. Jt. Anim., Mamm. 

p. 13, no. 183 (1842: Ualan) ; Jentink, Cat. Ost. Mamm. 

p. 2t)2 (.specimen f^) (1887: Ualan); id., Cat. Syst. Mamm. 

p. 149 (18'88 : Ualan). 
Pteropus keraudrenius (pt., ner Temm. 1825), Temminch, Man. 

Mamm. ii. p. 77 (1837: Ualan); Fitzinger, SB. Akad. Wien, 

Ix. .\bth. i. p. 437 (1870: Caroline!:). 
Pteropus ualensis, Finsch, Ibis, 1881, p. 107 (ITiilan) (ncn. nud.). 
I'teropus ualanus, Peters, SB. Ges. nat. Fr. 1883, no. 1 (Id Jan.) 

p. 1 (Ualau) ; Jentink, Cat. Syst. Mamm. p. 149 (1888 : Ualan? ; 

Tronessart, Cat. Mamm. i. p. 78 (1897 : Carolines) ; ? Ellint. Cat. 

Mamm. Field Col. Mus. p. 492 ( 1907 : Carolines) : (^. M. Allen, 

IhUl. Mug. Comp. Zool. Han: QM. lii. no.3,p. 29 (1908: Ualan). 
Pteropus (Spectrum) ualanus, Matsdiie, Meqachir. p. 28, pi. v. ligs. 

17-22 (; .skull) (1899: Uaiau); Troiu:<sait, Cat. .Mamm.. Supjd. 

p. 53 (1904 : Ualan). 

Diagnosis. — Similar to Pt. yapensis, hut with much smaller 
teeth. Porearm 130'5-133-5 mm. Ual>. E. Carolines. 


ISJcidl and iectli. — Skull as in Pi. yapenslx. Cheek-teoth on the 
whole much shorter (antcro-posteriorly) thon in Pt. yapcnsis, hut 
verj' nearlj' of the same width ; lonj^th of p'' 3'S— i mm. (against 4*5 
in the allied species), of p' 3-8-4 (against 4-3-4-5), of m' 4-2-4-5 
(.5-5-2), of m^ 4-4-1 (4-4-4-5), of m^ 3-3-2 (3-5-3-7). Structure of 
teeth as in Ft. yapensis. 

Colour. — Series, cotypes of species : Back and rump blackish, 
slightly sprinkled with silvery greyish hairs. — Underparts blackish 
seal-brown, siitiilnrly sprinkled. Foreneck sometimes slightly 
washed with dark russet. — Mantle, sides of neck, and occiput (as 
far as back of ears) huffy, pure in tinge or more or less clouded 
Avith russet ; base of hairs dark brown everywhere (females) or 
only in posterior ])ortion of mantle and on occiput (males). — Crown 
approaching vandyck-brown, with or without a slight wash of 
russet, gradually darkening on sides of head into the blackish 
seal-brown of chin and throat. 
-- Measurements. On pp. 182, 183. 
" i^liecimens e.ramined. Ten, cotyjies of species. 

Range. Eastern Carolines : Ualan (Kushai). 

Cot]ipes in the Berlin Museum. 

Pteropus nalanns, Peters; 1883. — Based on ten specimens 
obtained by Dr. 0. Finseh in Ualan, viz., mounted specimens 5798, 
5808, 5809, with skulls separate, and alcoholic specimens 5788 
(two), 5791 (two), 5792 (two), and ?5799 (one). Seven specimens 
are more or less young, two young adults (apparently full-grown), 
one adult. Skull of cotype 5808 figured in • Megachiroptera des 
Berliner Museums ' (7. s. c). 

llemarls. — This species is the Ualan re]ncsentativp of Pt. 
yapensis. It is scarcely distinguishable, with certainty, from this 
latter by any other character than the much smaller size of the 

20. Pteropns niariannus, Desm. 
Pteropus I'eraudreni (pt.), Dobson, Cat. Chir. B. ^1. p. C3. 

Pteropus mariannns, Deamarest, Mainm. ii., Suppl. p. ;j-17, footnote 

(1822: Mariannes). 
Pteropus marianus, Gi'ehel, Sttuy. p. 998 (1855 : Mariani es). 
Pteropus (Spectrum ) marianniis, Matschie, Meyuchir. p. 27, pi. v. 

tigs. 15, 16 (skull) (1809: Guam); Trouessarf, Cat. Mamni., 

(S'!/p^;/. p. 53 (pt.) (1904: Mariannes). 
Pteropus keraudren, Quoy ^- Caimanl, Voy. ' Uranir,^ Zool. pt. 2, 

p. 51, pi. iii. fig. ] (animal), fig. 2 (skull) (1824: Guain;; 

Lesson, Mail. Mamm. p. lU!), no. 281 (pt.) (1827); Bcsmarest, 

Diet. Sci. Nat. xlvi. ]). 364 (1827 : Mariannes) ; Is. Gvoffroy, 

Diet. Class, crilist. Nat. xiv. p. 701 (1828: Mariannes); J. B. 

Fisclier, Syn. Mamm. p. 84, no. 10 (1829 : Guam) ; Lesson, Hist. 

Nat. Mamm. {Compl. Bufon) v. p. 56 (1836: Mariannes); Oken, 

■ .1%. i\'«<?/r/-/Mc;(. vii. Abth. ii. p. 990 (1838); Le.oson, N. TaU. 

''■''■ i?. Anivi.,' Mamm. p. 13, no. 183 (pt.) (1812: Mariannes); 

E. Desmurest, Diet. Unir. d'llist. Nat. xi. p. 218 (1848 : Guam) : 

pti;roi'L's makianxl's. 179 

Ji'titink, Cut. Ost. Mamm,. p. 26:i, specimen 6.(1:887: (Tiiam); 

id.. Cat. Sijst. Mamm. p. 149 (pt.) (ISSH:- Gujim) ; Miller, Finn. 

Sf Gen. Bats, p. 58 (pt.) (1907). - 
Pteropus keraudrenius, Temminck, Man. Mamm. i. p. 186, pi. xv. 

fig. 7 (skull, copy from Qiioy & Gaimard) (1825: Guam) ; id.., 

op. cit. ii. p. 77 (pt.) (1837 : (juam) ; Wapier, Schrehcrs Saic;., 

Suppl. i. p. 35.3 (1839: Mariannes) ; Schinz, Si/st. Verz. Siiug. 

i. ]). 12(5 (1844: Mariannes); Wagner, 8chreher's Silug., Suppl. 

V. p. GOO (1853-55: Mariannes); Fitzingcr, SB. Akad. Wieii, 

l.\. Abth. i. p. 437 (pt.) (1870: Guam). 
Pteropus Iceraudreni, Grny, May. Zool. >.V But. ii. p. 503 (1838: 

(Juam) ; Gervai.->, Hist. Xat. ^[((mnl. i. p. 18'^, c. lig. (head) 

(1854: Guam); Peters, MB. Akad. Berlin, 1867, p. 331 (pt.) ; 

Dobsun, Cat. Cliir. B. M. p. 63 (pt.) (1878: Mariannes); 

Trouessart, liev. l^- Maij. Zool. (3) vi. p. 203 (pt.) (1879: Samoa, 

errore) ; Elera, Cat. Sist. Faun, Fili/iinas, p. 6 (pt.) (1895 : 

Mariannes) ; Omtalef, N. Arch. Ma.i. d'Hist. Nat. (3) vii. p. 14i» 

(1895: Guam; Rota; Saypan) ; Trouessart, Cat. Matiun. i. 

p. 82 (pt.) (1897 : Mariannes). 
Pteropus keraudreusis, Oketi, Ailtj. Xatunjeseh. vii. Abth. ii. p. 990 

Roussette Keraudren, Qnoij S,- Gaimard, Ann. Sci. Xat. vi. p. 111! 

(1825: Guam; " la petite ile aux Cocos ") ; Hd., J'oi/. ^Astrola/ir,' 

Zool. i. p. 80, pi. X. (1830 : anatomy). 

Diagnosis. — Similar to Pt. yapensis, but slightly larger and with 
considerably heavier dentition. Forearm 184-13()'5 mm. H(i/>. 

Skull and teeth. — General characters of skull as in Pt. yapcnsis, 
but size slightly larger. Dentition on the whole decidedly heavier, 
the increase in size being especially noticeable in p', p', and all 
lower cheek-teeth except p^ and m^.; see measurements, p. 183. 

Colour. — Cotype no. 50, J ad., Guam, and twelve specimens 
collected by Marche in the islands of Guam, liota, and Saypan 
(Paris Museum). — Back and rump blackish seal-brown, slightly 
sprinkled with silvery greyish-white hairs. — Breast, belly, and 
Hanks seal-brown (in eotype faded to chocolate), slightly mixed 
with silvery hairs. — Mantle and sides of neck ochraceous-buff ; 
foreneck often as pale as sides of neck, but in some specimens more 
or less hea\ily clouded with dark brownish. — Crown dark brown 
mixed with buffy hairs, the dark element becoming more pre- 
dominant on sides of head, and darkened to seal-brown on chin and 

Measurements. On pp. 182, 183. 

Specimens e.vamimd. Fifteen, in the collections of the Paris 
(fourteen : Guam, llota, Saypan) and Berlin Museums (one : Guam), 
including two cotypes of Pf. mariannvn and Pt. keraudren. 

Raufje. Marianne Island.? : Guam, llota, Saypan. 

Cotypes in the Paris and Leyden Museums. 

Pteropus mai-ianmis, Desmarest ; 1822. — Type locality, Guam 
(voyage of the ' Uranie ') ; cotypes the same as those of Pt. 
keraudren, Q. & G. 

J'teropus keraudren. Quov &. Gaiward ; 1*^24. — Based on four 


iSd i'ii;r.(ii'is .■\iAKiAXxrs. 

specimens, obtained in (mam during the voyage of tlie ' Uranie,' 
being the same series briefly diagnosed and named Pt. mariannus 
two years before by Desmarest. Tw') of these cotj'pes are in the 
Paris Museum, an adult female (A. 49, old catalogue 751) and an 
adult male (A. 50, eld catalogue 753 A), both mounted and some- 
what faded ; skull of A. 4U extracted in 1907 ; the skull of A. 50, 
which is not in the skin, is probably skull no. 0765 of the Museum 
d'anatomie comparee, which is marked " Mariannes, Quoy & 
(iairaard, Uranie.*' The two other cotypes were given in exchange 
to the Leydcn Museum ver}' soon after the arrival of the ' Uranie ' 
collection (see Temminck, Mon. Mamm. i. pp. 186-187, 1825, and 
Oustalet, 7. .s. c. 1895) ; they are the mounted specimens h and d. 
and skull b (of specimen d), of Jontink's Catalogues. Temminck's 
'■'Pt. l-eraudrcnius," of 1825 {I. s. c), is a rede.scription of these 
cotypes, accompanied by a copy of Quoy and Gaimard's figure of 
the skull ; in 183" (Mon. Mamm. ii. pp. 77-78) Temminck mentions 
two additional specimens of " Pt. keraudrenius "' collected by 
Ivittlitz and acquired by the Leyden Museum ; the one, from Guam, 
is Pt. rn((rianm<s, but the other, li-om Ualan. is I't. ualanus. — 
A topotype collected by Kittlitz is in the Berlin Museum (no. 344, 
5 juv., not full-grown) ; the skull of this specimen is the original 
of pi. V. figs. 15, 16 of the ' Megachiroptera des Berliner Museums.' 
Remarhs. — This species can apparently only be discriminated, 
with certainty, from the allied Pacific forms by the characters of 
the skull and teeth. In the size of the orbits (diameter 11'8 mm.) 
it accords with the N. Pacific Pt. yapensif. valanvs, and loochoensi.^ 
(11-5-11-8), and dift'ers from the h>. Pacific Pt. vanilorensis and 
ioncjanus (12-5-13-2). The dentition is on the whole heavier than 
iu the other N. Pacific forms, and as heavy as in Pt. vanilcorensis 
and tomjanvs. The external dimensions (forearm 134-1 36"5 mm.) 
may average slightly larger than in Pt. yapensift and ualanus 
(130-13.3"5), of whicli, however, only a few adult specimens have 
been available for comparison, and are an])ai-ently equal to those 
of Pt. loochonisis and tY«»7i-o>YMm (1 35-142-5). In some specimens 
of Pt. mariannv.o, including the Paris cotypes and tlie Berlin topo- 
type, the pale yellowish colour of the mantle and sides of neck 
extends over the whole of the forcneck, forming a complete collar, 
while in some Paris topotypes collected by Marche the collar is 
interrupted in front by a more or less considerable admixture of 
dark brownish in the centre of the foreneck (as is constantly the 
case in Pt. vanihorerisis and tQmjnnvs); the foreneck of /^/. mariannus 
no doubt averages paler (more unmixed yellowish buff) than in the 
!S. Pacific forms, but the character, as being in all species of this 
group subject to a good deal of individual variation, is of little 
practiciil use for diagnostic puri)Oses. 

rxKiiopL-s r.oociiot:xsis. 181 

"21. Pteropus loochoensis, Graif. 
Pteropiis l-eraudreni (pt.)) Dobson, Cat. Chir. B. M. p. 63. 

Pteropus loochoensis, Grai/, Cat. Monk. Sj-c. p. lOG (1870 : Liu-kiu 
Is.) ; 3Iiller, Fam. c<r Gen. Bats, p. oS • (1907). 

Pselapiion luchiien.sis, Seitz, 31iltli. I). Ges. Nnfurk. Osfasiois, v. 
p. 364 (1892: l.iii-kiiis). 

Pteropus keraudreiii, var. loochooensis, Fritze, Zool. Jahrb., Si/st. vii. 
p. 854 (1894: Okinawa). 

Pteropus (Spectrum) loochoensis, Matschte, Mer/acfiir. p. 27 (1899: 
Liu-kius) ; Troiiessart , Cat. Mamm., liupjil. p. 53 (1904 : 

Pteropus keraudreni loochooensis, L'onhote, Xm: Zool. ix. p. 628 
(1902: Liu-kius). 

Pteropus keraudreni {nee Pt. keraudren, Q. H; G.), var. /3, Dohson, 
Cat. Chir. B. M. p. 65 (1878: Liu-kius). Var. hh. Trouefsart, 
Rev. ^- Moff. Zool. (3) vi. p. 20:5 (1870: Liu-kius). Var. a, 
Trouessarf, Cat. Mamm. i. p. 83 (1897: Liu-kius). 

Dltifpiosis. — Similar to Pt. mariannus, but with smaller premolars 
and conspicuously longer fur. Forearm 135-1 42';'i mm. Ilab. Liu- 
kiu Is. 

Skull and teeth. — Skull as in Pt. mariunnus. Premolars abovo 
and below larger than in Pt. yapensis, but smaller than in Pt. 
mariannus ; see measurements, p. 183. 

Fur. — Longer than in the allied species; approximate length, 
back 14-17, mantle and belly I0-I8 ram.: com\rdrQ Pt. yapenslr 
■dxid j)elewensis. Least width of furred area of back about 42 mm. 

Colour (type, J yg. ad.). — Back and rump glossy blackish seal- 
brown thinly sprinkled all over with greyish-white. — Breast, bell}', 
and tlauks glossy scal-browu sprinkled with silvery greyish and 
buffy hairs. — Mantle bufl'y, palest (nearly cream-buff) posteriorly, 
tinged witli orange ochraceous in middle, and passing into ocbra- 
ceous-buff on sides of neck ; median line of foreneck much clouded 
with mars-brown and seal-brown. Concealed seal-brown bases of 
buffy hairs very short in centre of mantle, somewhat longer pos- 
teriorly in shoulder region, wanting on sides of neck. — Occiput 
similar to mantle or somewhat tinged with tawny ; crown mottled 
dark brown, russet, and buffy ; circumocular region conspicuously 
paler (owing to predominance of buffy element) than centre of 
crown ; cheeks seal-brown sprinkled with bufly ; chin and throat 
blackish seal-brown with some trace of paler admi.x.ture. 

A second specimen ( $ ad., teeth somewhat worn, ; 
Gray's "variety" of Pt. loochoensis, 1. s. c. p. 106) is lighter in 
colour, chiefly owing to a stronger admixture of whitish-grey and 
buffy hairs on back and underparts and a considerably paler colour 
of mantle and head. — Back blackish heavily mixed with buffy and 
greyish along median line, much less so on sides of back and rumj). 
— Breast. l)elly, and Hanks a little paler than seal-brown, heavily 
mixed with light greyish on sides of breast and belly, and with 
bufl'y on centre of breast; anal region slightly washed with mavs- 



browi). — Mantlo cream-buffy, tinged with buffy in centre and on 
sides of neck ; foreneck somewbat clouded with pale russet. Fur 
of mantle, sides of neck, and foreneck with rather long seal-bvoAvn 
bases (probably sesnal character). — Occiput similar to mantle ; 
crown and circumociilar region slightly darker: temi)oral region, 
chin, and throat seal-brown sprinkled with pale buflfy. 

Measurements. Below and on p. 183. 

Sjiechnetts e.vam'uied. Two, as registered below. 

Hange. Liu-kiu Islands (Okinawa). 

Ti/j'ie in collection. 

Beiiuirls. — This species is readily discriminated from Pf. mari- 
rinnus by the characters given in the diagnosis above. From 
J't. ionf/anns it differs chiefly in its weaker dentition, smaller skull 
with relatively smaller orbits, and longer fur ; the external 
dimensions avcraa;e smaller. 

2 ad. sk. ; skull. 
c? yg. nd. sk. : .skull 

Liu-kin Is. 
Liu-kiu Is. 

Purchased (Warnick). 4'ALo.l. 
Purcbased (Warwick). < 401 "ifi''< 

{Type of species.) 

External measurements of Ptpropus ualanns, roariannus, 
and loochoensis. 


Polles, total length, c. u 

,, metacarpal 

,, 1st phalanx 

2iicl digit, metacarpal 

,, Lst phalanx 

„ 2nd-3rd phalanx, c. u 

3rd digit, metacarpal 

,, 1st phalanx 

,, 2nd phalanx 

Ith digit, metacarpal 

1st phalanx 

, . 2nd phalanx 

•3th digit, metacarpal 

,, Ist phalanx 

2Rd phalanx 

Ears, length from oi-ifice 

,, greatest vidth, flattened 
Front of eye to tip of muzzle .... 

Interf emoral 

Lower leg 

Foot, c. u 


* Reg. noa. 5788 (2 cT yg. ad. al.), 5808 ( c? id. mounted) ; Berlin Museum. 
t Eeg. nos. A. 49 ( 2 ad. mounted), A. 50 (c? ad. mounted) ; Paris Museum, 
t Estimate (skins). 

PTimorrs ttalvxts etc. 


Measurements oj shulls and teeth of Pt. ualaiius, mariamuis, 
and loochoensis. 

Pt. wflanus. 

Pt. marianniis. 

Pt. loocho'eusif. 

Skull : 1 ad. 


: 2 ad.t 


Teeth : 1 ad., 

Teeth : 

2 ad.t. 



2 iiniu. 

1 imm.J 





5 ad. 

Mix. JIa.v. 




mm. mm. 






liU, total leugtli to gua^hioii 





, palation lo incisive foramina ... 

trout of orbit to tip of nasals... 






width of braiu-case at zygomata. 




, zygomatic width 






witllh across m', externally 

hiclirvuiiil width 






width across canines, externally 

, postorbitjl constriction 








mterorbital constriction 

width of iiiesoptcrvgoid fossa... 

between p''-p'', internally 






between cingula of canines 













iiidible, length 

„ coroiioid heijjht 





Upjjer teeth, c-m^ 


24 5 










)per incisors, combined width 



3-8 4 
31 3-5 




4-8 1 
33 1 




3-8 4 





3-2 3-4 
4-2 4 5 
2-7 2-8 
1-8 2-2 










, length 





, length 



1-2 1-8 








1-7 1-8 




1-8 19 
4-2 4-5 









2-8 3 




3 ; 


4 41 

31 3-2 



3 2 





, length 

4 41 , 






28 29 






, length 

3 3-2 






2-2 2-5 






, length 

1-5 1-6 







1-5 1-5 




2 ! 

* Reg. no8. 5808 (cC ad.), 5798 (2 hnm.). 581)9 (J imra.); Berlin Museum. 

t llcg. nos. A. 49 (eotype), A. (i7(J5 (^topotype, collected by Marche) ; Paris Museum. 

t Keg. no. 344 (topotype, collected by Kitllitz); ]3erlin Museum. 

18-t rTEUons taxikohexsis. 

22. Pteropus vanikorensis, Q. c|- G. 
Pteropns kei-audreni (pt.), Dobsoii, Cat. Ohir. B. M. p. 63. 

r Pteropus cliissuniieri, Is. Geoffroy, Diet. Class. tVHist. Nat. xiv. 
p. 701 (1828 : India ; Amboina) ; J. B. Fischer, Syn. Mamm. 
p. 85, no. 11 (1829 : India ; Amboina) ; /*•. Geoffroy, in Belanyers 
Voy. Indes Orient., Zool. p. 98 (1834 : India; Amboina) ; Lesson, 
Hist. Nat. Mamm. (Com})!. Bvffou) v. p. 48 (1836: India; 
Amboina): Teinniinck, Mon. Mamm. ii. p. 76 (1837: Indian 
continent; not Amboina); Gray, Mag. Zool. Sc Bot. ii. p. 503 
(1838: India; Amboina); Warner, Schrelter's Siiug., Suppl. i. 
p. 355(1839: India; Amboina); Ogiiby, Madras Journ. Lit. S,- 
Sci. xii. p. 146 (1840) ; Lesson, N. Tahl. R. Anim., Mamm.^. 13, 
no. 181 (India); Schinz, Syst. Verz. Sang. i. p. 127 (1844: 
Indian Peninsula) ; E. Desmarest, Diet. Univ. d'Hist. Nat. xi. 
p. 248 (1848: India; Amboina); Wagner, Sckrebers Siiug., 
Suppl. V. p. 601 (1853-55: ? Indian continent; P Amboina); 
Gervais, Hist. Nat. Mamm. i. p. 188 (1854 : India) ; Giebel, 
.S'rtvy. p. 988, footnote (1855: India; Amboina); Fitzinger, SB, 
Akad. Wien, Ix. Abtb. i. p. 442 (1870 : Indian Peninsula). 

Pteropus vanikorensis, Quay iS'- Gaimard, Voy. ' Astrolabe,' Zool. i. 
p. 77 (specimens, not skull), pi. ix. (animal) (1830: Vanikoro) ; 
■r Lesson, Hist. Nat. Mamm. (Compl. Buffon) v. p. 58 (1836: 

Vanikoro) ; TemmincTt, Mon. Mamm. ii. p. 78 (specimens, not 
skull) (1837: Vanikoro); Wagner, Schreber's Saug., Suppl. i. 
p. 354 (1839 : Vanikoro) ; E. Desmarest, Diet. Univ. d'Hist. 
Nat. xi. p. 250 (1848 : Vanikoro) ; Gervais, Hist. Nat. Mamtn. 
J. p. 188, c. fig. (head) (1854 : Vanikoro); Giebel, Siiug. p. 998, 
footnote (1855 : Vanikoro). 

Pteropus Tanicorensis, Oken, AlU). Naturg. vii. Abth. ii. p. 990 
(1838); Schinz. Syst. Verz. Siiug. i. p. 127 (1844: Vanikoro): 
Wagner, Sc/ireber's Scim/., Suppl. v. p. 601 (1853-55 : Vanikoro) ; 
Fitzinger, SB. Akad. Wien, Ix. Abth. i. p. 441 (1870: Vanikoro). 

Acerodon vanikorensis (pt.), Lesson, N. Tabl. Ii. Anim., Mamm. 
p. 14, no. 194 (1842 : Vanikoro). 

Pteropus keraudreni (pt., nee Pt. keraudren, Q. 4- G.), Peters, MB. 
Akad. Berlin, 1867, p. 331 ; Dob.wn, Cat. Cliir. B. M. p. 63 (1878) ; 
Trouessart, Rev. <§• Mag. Zool. (3) vi. p. 203 (1879 : Vanikoro) : 
id., Cat. Mamm. i. p. 82 (1897: Vanikoro). 

Pteropns tonganus (pt., 7ieo Q. Sj- G.), Matschie, Megachir. p. 19 
(1899 : Vanikoro) ; Trouessart, Cat. Mamm., Stippl. p. 51 (1904 : 

Diagnosis. — Similar to Pt. mariannns, but with slightly larger 
skull and relatively much larger orbits. Forearm about 186-137 
mm. Hah. Sta. Cruz Is. 

Skull and teeth. — Skull averaging rather heavier than in Pt. 
mariannvs and loocJioi'nsis ; orbital diameter 13 mm., against 11-5- 
ll'S in the allied species. 

Fur. — As in Pt. fo))ija)ius. 

Colour. — Cotyjie. J ;(d.(Kog. no. A. 58). — Buck and rump dark 
?eal-brown, thinly sprinkled Avith greyish-white hairs. — Breast, 
bcllv. Mild flanks similar, but flanks nither darker th;in cpidre of 


breast and belly. - Mantle ocliraceous-buff, lighteninf^ to buffy or 
crcam-bufty posteriorly ; occiput and sides of neck similar to 
mantle, but rather more tinged with ochraceons ; I'lir of foreneck 
ochraceous at base, seal-brown at tip, making colour of foreneck 
superficially like that of breast. — Forehead, crown as far back as 
front of ears,, sides of head, chin, and throat seal-brown. 

Cotype A. 57 (J ad.) is in poor condition, but has apparently 
not differed in any noteworthy respect from foregoing. 

Measurements. On pp. liJO, 191. 

Sjjecmens examined. Four, in the collection of the Paris Museum, 
including the two cotypes of the species and one cotype of 
Ft. rlussumieri. 

Range. The island of Vanikoro. 

Cotypes in the Paris Museum. 

Pteropvs duss%(mierl,ls. (jQo^ve^y ; 1828. — Based on two speci- 
mens, the one stated to have been brought by Dussumier from the 
continent of India, the other supposed to be from Araboina. The 
former is in the Paris Museum (Reg. no. A. 61 ; 2 ad. mounted, 
skull in situ) ; it has been acquired through Yerreaux and is 
ticketed "Bengale?": the latter I have not been able to find in 
the Paris collection, nor iu the manuscript register of Chiroptera of 
that Museum. Both of the localities given by Is. Geoff'roy are 
undoubtedly incorrect. The cotype examined by me is externally 
indistinguishable from Pf. vanikorensis, but the characters of the 
skull and teeth are in this group of the genus of such decisive 
importance that the identification of Ft. dussumieri remains 
uncertain so long as its skull has not been made available for 

Fteropns vanil-orenfiis, Quoy and Gaimard ; 1830. — Original 
description based on two specimens obtained in Vanikoro during 
the voyage of the ' Astrolabe ' (Paris Museum, E,eg. nos. A. 57 and 
A. 58), both males, mounted, skull of A. 58 in situ, of A. 57 
extracted in 1907 ; cotype A. 58 is probably the original of Quoy 
and Gaimard's pi. ix. A third mounted specimen in the Paris 
collection (A. 59, not full-grown), though marked "P. vanilor- 
eiisis, Q. & G. ; type ; Vanikoro : Quoy & Gaimard, Astrolabe," 
cannot technicallj' be considered a cotype, since it is not referred 
to in the original description. — The skull described by Quoy and 
Gaimard as being that of Ft. vanilcorensis (Paris Museum, no. 6746) 
belongs to a totally diflerent species, Pt. tuhereidatus, Peters. 

lientarh^. — This species is readily distinguished from Pt. mari- 
annvs by its lai'ger orbits. From the closely allied Pt. tonganus 
it differs by its slightlv smaller size (forearm 136-137, against 
139-150 mm.) and relatively shorter metacarpals (third metacarpal 
88-5, against 93-5-1 02-5 mm.). 

1S6 nintorrs toxgam's. 

'SS. Pteropiis tonganus, Q. 4- G. 
Ficrojms keraudreni (pt), Dobson, Cat. Cliir. E. M. j). 63. 

I'teropus ton^'^anus, Quoy cS' Gaimard, Voij. ' Astrolabe,^ Zool. i. p. 74, 
pi. viii. fig'. 1 (col. tio-. of young), figs. 2, 3 (skull), lig. 4 (col. tig. 
of pale variety, young) (1830: Tonga-Tabu); Lesson, Hist. 
Nat. Mamm. \Cvmpl. Buff'on) v, p. 57 (1836: Tonga-Tabu); 
Temminck, Mon. Mamm. ii. p. 79 (1837 : Tonga-Tabu) ; Oken, 
Allg. Natnrq. vii. Abth. ii. p. 990 (1838); Wayner, Schreber's 
Sciitg., Suppi. i. p. 353 (18-39 : Tonga-Tabu); Lesson, N. Tahl. B. 
Anim., Mamm. p. 13, no. 184 (1842 : Tonga); Srhinz, Syst. Verz. 
Sauy. i. p. 120 (1844 : Tonga-Tabii) ; U. IJe-wwresf, Diet. Unit-. 
d'Hist. Nat. x'l. p. 248 (1848 : Tonga) ; Wayner, 8c/ireber's Sauy., 
Siippl. V. p. 60() (1853-55: Tonga-Tabu); Gevvais, Hist. Nat. 
Mamm. i. p. 188 (1854: Tonga) ; Giebel, Siiuy. p. v;)98, footnote 
(1855: Tonga): Fitzinyer, SB. Akad. Wien, Ix. Abth. i. p. 439 
(1870: Tonga-Tabu); Marchi, Atti Soc. Itul. Sci. Nut. xv. 
p. 516 (1872-73: structure of hairs) ; Gibdher, P. Z. S. 1874, 
p. 295 (Savage I.) ; Mutschie, Meyachir. p. 19 (pt.) (1899 : Tonga ; 
Wallis Is.; fSanioa) ; Troucssart, Cat. Mamm., Svppl. p. 51 (pt.) 
V (1904: iMJi ; Tonga; Wallis Is.; Samoa). 

ji Pteropus keraudrenius [nee Temm.), Peale, U.S. E.vpl. Exp. viii., 
■) Mamm. p. 18 (1848: Fiji); Cassin, op. cit. viii. (sec. ed.) p. 10 

(1858: Fiji). 

Pteropus keraudreni [7iec Pt. keraudren, Q. Sf G.), Schmeltz, Cut. iii. 
DoM. Godeffroy, p. 1 (1866: Fiji): Peters, MB. Akad. Berlin, 
18H7, p. 331 (pt.) ; Dobson, Cat. CMr. B. M. p. (33 (pt.), p. 552 
(1878: Fiji; Samoa; Savage I.) ; Troiiessart, Rev. Sf May. Zool. 
(3) vi. p. 203 (pt.) (1879: Fiji; Samoa; Savage I.); Moseley, 
Notes Naturalist ' Challenyer,' p. 291 (1879 : Tonga-Tabu ; habits) ; 
'Troue.'sart, Ann. Sri. Nat. (6) Znol. viii. Art. 12, pp. (j, 17 (1879 : 
remarks on distribution) ; J. Anderson, Cat. Mamm. Ind. Mus. 
i. p. 103 (1831: Samoa); Lydekker, Li. Nat. Hist. i. p. 256 
(1893-94 : Tonga) ; Trouessart, Cat. Mamm. i. p. 82 (pt.) 
(1897 : Fiji ; Tonga ; Samoa). 

? Pteropus funereus, Kreffl, Cat. Mamm. Austral. Mus. p. 4 (1864: 

Pteropus flavicollis, Gray, Cat. Monk. S,c. p. 107 (1870 : Moala ; 
Totoya ; Ovalau) ; Alston, P. Z. S. 1874, p. 96 (Samoa). 

Diagnosis. — Similar to Pt. ranikorensis, but rather larger and 
■with relatively longer wings. Forearm 139-150 mm. JJah. 
Fiji, Tonga, and Samoa Is. 

Differential characters. — This species is scarcely distinguisliable 
from Ft. vaniJiVroisis by other characters than those given in tho 
diagnosis : forearm 139-150 mm., against 136-137 mm. in the 
cotypts of l^t. vanilcorensis ; third metacarpal 93-5-102-5 ram., 
against 8S"5 ; compare measurements p. 190. Size of orbits 
( t 2-5-13-2 mm.) as in Pt. vanil-orensis (13 mm.). 

Palaie-ridqcs. — 54-5-f-3, but not rarely with a more or less 
incomplete additional ridge between the normal ninth and tenth 
(formula approacliing 5-|-6-f3). Arrangement of ridges as in 
Pt. liypomclanus (above p. 101). 

Fur. — Longest hairs of back approximately 10-13 mm., of mantle 

rtEuorvs toxganits. 1S7 

12-13 in malfs, l(i-17 in females, of bdly iy-15. Dibliibulioii of 
fur as ill the allied species. 

Cohvv. — British Museum series, males and females, adults and 
iinraatures (see list below): — Back, rump, breast, belly, and flanks 
blackish or seal-brown, generally slightly darker and more glossy 
on back and rump than on breast and belly, everywhere thinly 
sjjrinkled with whitish hairs.- — Mantle l)uffy, sometimes very pale 
(approaching cream-buff), sometimes distinctly washed with oehra- 
ceous-buff or pale orauge-ochraceous ; sides of neck similar or 
slightly deeper in tinge. Base of fur of mantle and sides of neck 
varying in colour according to sex of individuals (see below). 
Foreneck ochraceous-buff)- more or less strongly clouded with pale 
russet and blackish seal-brown ; in many specimens the buffy 
colour is purest (less mixed with darker tinges) on posterior portion 
of foreneck, producing a more or less distinct narrow transverse band 
separating dark breast from dark centre of foreneck ; in other 
specimens the dark admixture extends over the whole of the lore- 
neck. — Occiput, as far as a line between middle or front of ears, 
similar to mantle or slightly mixed with brownish. Crown mottled 
dark brown and bufty, in some specimens nearly uniform seal-brown 
like back. Temporal region and cheeks similar to crown or more 
uniform dark brown ; circumocular space in dark-headed s])e- 
cimens often more or less strongly suffused with pale russet or buffy 
russet, forming more or less distinct " spectacles " (cf. Pt. coiispi- 
cillatus). Chin and throat blackish or seal-brown. 

Sexual differentiation.- — Hairs of mantle in adult males shorter, 
more rigid and oily, in adult females longer, softer, more spreading. 
In all males examined the hairs of the mantle and sides of neck are 
uniform buS'y to extreme base, or with short blackish bases only 
posteriorly in shoulder region : in females the fur of the mantle 
and sides of neck has long concealed seal-brown bases. 

Mteisuremcnts. On pp. 190, 191. 

Specimens examined. Twenty-two, in the collections of the Paris 
(three), U.S. Xational (two*), and British Museums, including the 
cotypes of tlic species and of Ft.jiavieoJJis. 

liani/e. Fiji Is. (Taviuni, Ovalau, iloala, Totoya) ; Tonga 1«. 
(Uea, Namouka, Tonga Tabu); Samoa Is. (Savage I.). The Tonga 
and Samoa groups form the extreme south-eastern limits of the 
range of the genus in the Polynesian region. 

Cofi/pe.<i in the Paris Museum. 

Habits. — Like many other species of the genus, Pt. tonga nus 
appears sometimes on the wing in the early afternoon in full sun- 
light, but as dusk comes on it becomes more and more plentiful, 
and it is probably only those accidentally disturbed or specially 
driven by hunger that come out before dusk. At the time of the 
visit of the ' Challenger ' to Tonga Tabu (July, 1874) these Fruit- 
bats were feeding on the bright red flowers of one of the indigenous 
trees, and flowers would seem on the whole to form an important 
proportion of the food of the species of Ptcropns ; they probably 
* Xos. U-2i72, U-2\ir>, Xamouka, Ton-a Is. 

ISS r-naiurx's TONOAXvg. 

often act as fertilizers by carryin<i; polloii from tree to tree, adherent 
to their fur. (Moseley, I. s. c.) 

Pterojnis tonganus, Q,\\o\ & Gaimanl ; 1830. — Oriofinal descrip- 
tion based on three specimens, o!)tained iu Tonga Tabu (voyage of 
the 'Astrolabe'), viz. nos. A. 54, A. 55, and A. 56, all mounted 
with skulls in situ. Xo. 54 (old catalogue no. 754) is an adult 
male ; no. oo ("75) is a young female, no doubt the origiual of the 
coloured figure pi. viii. fig. 1 (drawing and coloration bad); no. 56 
(755 B), also young, is the yellowish individual figured pi. viii. fig. 4. 
Colour and other external characters of cotype 54 ( c? ad.) quite as 
in the British Museum series. Topotypes ('Challenger' Expedi- 
tion) are in the collection of the British Museum. 

Pteropits JlavicoUis, Gray; 1S70. — Based on specimens from 
Moala, Totoya, and Ovalau Islands, I'^ijis (voyage of the ' Herald '). 
Moala may be fixed as the type locality ; cotypes, c ad., 5 ad., 
skins with skulls, in collection. 

a. Ad. skull. Fijis(Voy.H.M.S. Musoiun of Economic 

■ Herald ' ; J. 

Geology [P.]. 



(S ad. al. 


^Museum Godeffroy. 


c. d. 

sks.; skulls. 

Taviuni, Fijis. 

£. A. Liardet, Esq. 


S ad., 2 ad. 

Ovalau, Fijis; 

Lords of the Admi- 


sks., skulls; 

Aug. 1856 

ralty [P.]. 

1 59.7.142. 

pull, skull. 

(Voy. H.M.S. 
' Herald ' ; Dr. 
F. M.Eai/7ier). 


[d] ad. sL; 

Moala, Fijis; 

Museum of Economic 


Sept. 1854 
(Voy. H.M.S. 
'Herald'; J. 


Geology [P.], 


$ nd. sk.; 

Moala ; Sept. 

Lords of tlie Admi- 


1854 (Voy. 

ralty [P.J. 


(li, i, cot.i/pe:< of Pt.JlavicolUs, Gray.) 


' Herald ' Dr. 

F. M. n.). 


Tull. sk. 

Moala ; riept. 
1854 (Voy. 

• Herald ' ; Dr. 
F. M. 11.). 

Lords of the Admi- 
ralty [P.]. 

Not reg. 


ad. fk. ; 

Totoya, Fijis ; 

Lords of tlie Admi- 


Aug. 1855 
(Vor. H.M.S. 
' Herald ' ; Dr. 

ralty [P.]. 

I, m. 

[r^l ad. sk., 
$ ad. al. ; 

Tonga Tabu 

Lords of the Treasury 

(Voy. H.M.S. 




' Challenger 'i 


Jnd., 2ad.. 
5 .iuv. al. ; 
.skulls of 


Museinu Godeffroy.,2,6 

9. '■• 

H, 0. 



Rev. J. S. Whit nice 



9 juv. nl.; 

Savage T.. 

Dr. A. Glint her [P.J, 



I'TEituiTs liF.niniir. ]S*J 

24. Pteropus geddiei, MurGil/iviui/. 
I'Uropus Ice rail dre id (pt.), Dohson, Cat. Chir. B. M. p. 0:5. 

Pteropus geddiei, MacGUUvray, Zoologist, x\ni. p. 7V-H (Sept. 1800: 
Aueiteum ; habits) ; Peters, MB. Akarl. JJerlin, 1867, p. 320 
(New Hebrides) ; Grai/, Cat. Monk. J^-c. p. 107 (1870 : Aneiteum) ; 
Matxchie, Mer/achir. p'l. vii. tigs. 3, 3tt, 3 b (skull) (1899: Now 

Pteropus keraudreni (nee Pt. keraudreii, Q. •^- G.), Dobsun, Cat. Chir. 
B. M. p. m (pt.) (1878 : Aneiteum) ; id., P. Z. S. 1878, p. 87o 
(1879: New Caledonia): Troiiesgarf, liev.'^- Ma(/. Zool. (3) vi.p.20-'> 
(pt.) (1879: AneiteuiuV; Dobson, Pep. Brit. Assoc. 1880, p. 17-'i 
(New Caledonia); TroveAsart, Cat. Manim. i. p. 82 (1897; 
Aneiteum; New Caledonia). 

Pteropus keraudreu (pt.), Jentink, Cat. Syst. Mamm. p. 149 (1888: 

Pteropus tongauus (pt., nee Q. S,- G.), Matxchie, Mec/achir. p. 10 
(1899 : New Caledonia ; New Hebrides) ; Troiie!i.''art, ( 'at. Maimn., 
Suppl. p. 51 (1904 : New Caledonia; New Hebrides). 

Diar/nosis. — Similar to Ft. tom/amis, hut sknlJ. teeth, and external 
dimensions larger. Forearm 153'5 mm. IJalt. New Hebrides ; 
New Caledonia. 

Colour. — Two skin.s, (J ad. (type). 9 ad. — Scarcely different 
from that of Pt. toncjrmns, except perhaps in the more uniform 
blackish colour of the bead. In both .specimens the crown and 
sides of liead arc as blackish as chin and throat, very sharply 
marked off from bufty occiput and mantle; rostrum and super- 
ciliaries distinctly Mashed with mars-brown or huffy mars-brown. 
The darkest-headed specimens of J't. foiir/anvs are, however, prac- 
tically indistinguishable in colour from tlie type of Pt. r/eddiei. 

tSexiud differentiation. — As in Pt. toni/anns. 

Measurements. On pp. lOl', 191. 

Specimens e.vamined. Four, in the coUectious oC I he Berlin (two, 
New Caledonia) and British Museums, including the type of the 

Jian(/e. Aneiteum; New Caledonia. 

Type in collection. 

Habits. — On a thickly wooded bank near the sea, on tiic leeward 
side of Aneiteum, behind a grove of bread-fruit and other trees, 
Pt. f/eddiei was found in great numbers by John MacCillivray, in 
1854 and ISoO, while for weeks and even months togetlier there 
were none in the neighbourhood. At this locality their arrival 
and departure were apparently connected with the ripening of the 
two annual crops of bread-fruit, to which they are very destructive. 
Jjike most other I'teropi, they are gregarious, but the sexes gene- 
rally keep apart; MacOillivray never shot any but males at the 
camp referred to above, nor did he ever meet with the female. 
They have their favourite trees for roosting on during the day. 
generally some wide-spreading fig or barivHn, sometimes an Acacia. 
During the day time and while suspended from tlie tree, they are 
readily roused by any unusual r.oise, such as the crackling of a stick 
under foot, and are jiot very easily approached. Like other large 
liats thry iiri' vrry tcn.ieious of life, and cling to the branches as 



long as they possess the requisite strength, and sometimes remain 
suspended even after death. The males seem to be very pugnacious ; 
they may be seen fighting among themselves on tho trees, and a 
wounded one will fui'iously attack anything brought in contact 
with it. AVith their formidable canines they inflict severe bites, 
and retain their hold with great tenacity. The Aneiteumese are 
fond of these bats as food, and spin the fur into cordage used in 
ornamenting the person. Besides occasionally killing them with 
stones and short sticks, they sometimes catch them in traps similar 
to those used for fishing— circular, flat-bottomed buckets made of 
FlageUaria inclica, with a hole at the top for entrance, and some 
papaw-apple for bait. The Aneiteumese generic name for Pteropus 
is Nekrei ; Pt. fjeddiei is called Xawathclgau, in contradistinction to 
Pt. anetianus, which is called Xalivatran, (ilacGillivray, I. s. e.) 

J. MacGillivrav, Kot reg. 
Esq. [C.]. 

{Ti/pe of species.) 
Canon H. B. 93. 11. -29.2. 

Tristram [E.]. 

External measurements of Pteropus vauikoreiisis, tonganus, geddiei. 

(I. c? ad. sk. ; 

/,. [2] ad. sk. 

Aneiteum, New Hebrides ; 
23 June, 1869. 


Pt. vaiiikoreiisis. 

3 ad. 
(Two cotypes of 
species, one of 
Pt. d'Ussumieri.) 

I Pt. tonganus. 

15 ad. 

(Incl. one 


Pt. geddiei. 

2 ad. 
(Incl. type.) 

I mm. 

Forearm ' 137 

Polles, total length, c. u 

,, metacarpal 14'5 

,, 1st phalanx SOo 

2nd digit, metacarpal 

„ 1st phalanx 

,. 2nd-3rd phalanx, c. u . ... 

3rd digit, metacarpal 

,, 1st phalanx 6G 

,, 2ud phalanx 

4tl! digit, metacarpal 

„ 1st phalanx 

2nd phalanx 

oth digit, metacarpal 

„ 1st phalanx 40 

2nd phalanx 40 

Eai's, length from orifice 

„ greatest width, flattened 

Front of eye to tip of muzzle 


Lower leg 62 

Foot, c. u 


* Cotype of Pt. vanikorensis (A. 57). 
|- Cotype of Pt. vamkorenMs' (A. 58). 















MiN. Max. Min. Max 






























76 5 































46 5 




i 'o§ 


^ .o0-5 
18-5 § 

20 § 

X Cotype of Pt. dttssir,niei-i {A.. 61). 
§ Estimate (skins). 



Measurements of sl-uJh and teeth 0/ Ptcropns vaulkoiensis, tonganns, 
and geddiei. 



Impedes.- — Pt. dohsoni, cnniceps, and argentatas. 

Itange. — North-western Anstro-Malaya : Celebes, Sanghir, Gilolo 
group, Sula Islands, ? Amboiim group. 

General characters. — In all essential respects similar to Ft. hypo- 
melanus and allies, but skull and dentition lieavier, ears generally- 
longer and distinctly attenuated above, colour of body characterized 
by a heavy sprinkling of dark fur with longer pale greyish or straw- 
yellow hairs, the pale element occasionally predominant, particularly 
on underside ; at least in one species a uniform mars-brown phase 
occurs. Sexual differeutiation inconspicuous (canines heavier iu 
males). Size below medium (forearm 135-145 mm.). 

Differentiation of species. — In the size and shape of the ears 
Vt. dohsoni (Celebes) comes nearest to Ft. Iiypomelanus, but the 
colour of the back is peculiar, sepia suflused with golden biiffy. In 
Ft. caniceps (Gilolo group, Sanghir, Sula Islands) the ears are 
longer, more attenuated above and narrowly rounded off at tip ; 
colour, in the ordinary grizzled phase, much paler. Ft. argentatas 
(?Amboina) is similar to i'?. «r/»'(r^j.s in the size and form of the 
ears, but much smaller-toothed, with longer and differently 
coloured fur. 

Affiaities of group. — Probably closely allied to the Ft. Jtj/po- 
nielaiius group, with which it accords in the general characters of 
the skull and teeth, distribution of fur, narrowness of central 
iiiterfomoral, and non-development of conspicuous secondary sexual 

2'). Pteropus dobsoni, K. And. 
Fteropus faacu^. Bobson, Cat. Chir. B. M. p. oD. 

I'teropus fuscus {nee E. Geoff , nee Dexm., nee Bkmiv.), Dobson, 

I. s. c. pp. 59, 552, pi. iv. tig. 5 (teeth) (Jane, 1878: Celebes); 

Trouessart, liev. ^- May. Zool. (3) vi. p. 202 (1879: Celebes); 

id., Cat. Mamm. i. p. 82 (1897 : Celebes) ; Miller, Fam. (!(- Gen. 

Bats, p. 58 (1907). 
Pteropus (Spectrum) liypomelanus /;. fuscus, Mntschie, Mcyaehir. 

p. 25 (1809). (/. fuscus, Trouessart, Cat. Mamtn.. Suppl. p. 52 

(1904: Celebes). 
Pteropus dobsoni, K. Andersen, Ann. S,- Mag. A*. II. (8) ii. p. 370 

(1 Oct. 1908). 

Diagnosis. — Similar to Ft. hypomelanus, but with broader 
rostrum, larger teeth, relatively larger p,, longer wings and tibia, 
and different colour of the fur. Back sepia suffused with golden 
buify ; sides of head and entire underside dark vandyck-brown or 
seal-brown ; mantle burnt-umber, strongly contrasting with back, 
but not with underparts. Forearm 144-5 mm. Hah. Celebes. 

Shdl. — Similar in characters and general size to that of the 
larger races of Pt. hypomeJamis, but rostrum slightly broader ; 
lachrymal width in type, a young adult male, 16 mm. (13-2-15-7 



iu all races of Pt. lirj pomelnnus) ; maxillary width externall}' across 
ni'-m' 19-5 mm. (16-5-19). 

Teeth. — Scarcely differing in structure from those of Ft. hypo- 
melaniis, but cheek-teeth conspicuously larger, particularly broader; 
m*, length (longitudinal diameter) 6-2 mm., against 4-S-5'9 in all 
forms oi Ft. hi/jiiomdaniis, -width. 3-7, against 2-9-3'5 ; m,, length 
5-8 mm., against ■4-2-5-2, width 3*S, against 2'8-3-2. The increase 
in the size of m^, as compared with Pt. hypomelanus, appears to be 
relatively larger than of the other cheek-teeth. For further details 
compare measurements, p. 109, with those of Ft. hypomelanus, 
pp. 1 18 and 131. — Cingulum of upper canines well defined but 
narrow. Posterior ledge of p' and p' short but distinctly marked 
off from tooth postero-externally. i^ fully twice the size of i,. 
p, about twice the size of i.^, larger than m^. Posterior ledge of 
Pj, and p^ quite short and rather ill-defined. — The peculiar shape 
and position of m" described and figured by Dobson {l.s.e.) is, 
no doubt, an individual abnormality in^tlie single specimen known. 
Ekirs. — As in Ft. hypomelanus ; scarcely attenuated above, tip 
rather narrowly rounded off. 

Wings and tibia. — Membranes arising about 15 mm. apart from 
sides of back. Length of forearm (144-5 mm.) in a slightly 
immature specimen as in the largest specimens of Ft. liypomeJanus 
(maximum 145'5 mm.), but metacarpals and phalanges decidedly 
longer ; compare measurements, p. 198, with those of Ft. hypo- 
melanus, p. 130. — -Lower leg G8o mm., against 54-5-66 in all races 
of Pt. hypomelanus. 

Inierfemoral. — Practically undeveloped in centre. 
Far. — Closely adprossed on back. Length, back about 11, 
mantle 13, belly 12 mm. Least width of furred area of back 
about 37 mm. Forearm and tibia naked above. 

Cohw (type). — Back and rump sepia, strongly and uniformlv 
suffused with a peculiar tinge of golden huffy approaching Isabella; 
seal-browQ base of fur concealed on back, partly exposed on pos- 
terior portion of rump. — Breast, belly, and flanks dark vandyck- 
brown thinly sprinkled with pale golden buffy hairs. — Mantle burnt- 
umber with mara-browu bases to the hairs, darkening to seal- 
brown with vandyck-brown hair-bases on sides of neck and 
foreneck. — Occiput and crown similar to mantle, passing into 
seal-brown on sides of head, throat, and chin. 
Measurements. On pp. 108, 199. 
Specimen examined. One, the type. 

Ilange. The single specimen on record is ticketed Celebes. 
Type in collection. 

Nomenclature. — The name proposed by Dobson for this species is 
preoccupied by Ft. fuscus, E. Geoff., 1803 { = Ft.niger, Kerr, 1792), 
Pt.fuscvs, Desm., 1803 {=Pt. stihniger, Kerr, 1792), and Ft. fuscvs, 
Blainville, 1840 (= Ft. vampyrus malaccensis, K. And., 1908). 

a. c? yg- afl- sk. ; skull. Cel«)ies. Purchased (Frank). 

( Type of sjieoies anil of P/./n.-^us, Dobson.) 

1'j4 PTEnorus caniceps, 

26. Pteropus cauiceps, (h-aii. 

Pteropus caniceps, Dobson, Cat. Chir. B. M. p. fiS. 
Pteroinis mackloti (pt.), Dobson, oj). cif. p. 07. 

rtei'opus hypomelanus (pt.), Temminch, Esq. Zool. p. G2 (1853: 

Ternate)'; Jentinh, Cat. Ost. Mamm. pp. 261-202, specimens o, 

j, l:,m, n (1887: Ternate; Gilolo ; Morotai ; Siao); id., Ccd. 

Syst. Mamm. pp. 148-149, specimens «, 6, e, /, m, o, p, s, t, u, v 

(i888: Ternate; Gilolo; Morotai; Siao; Sula Bessi). 
Pteropus caniceps, Gray, Cat. Monk. Sfc. p. 107 (1870 : Batchian) ; 

Dobson, Cat. Chir. B. M. p. 08 (1878: Batchian; Gilolo): 

Trouessart, Rev. ^- Mag. -Zooi. (3) vi. p. 203(1879: Batchian ; 

Gilolo); Jentink, Cat. Syst. Mamm. p. 150 (1888: Batchian); 

Trouessart, Cat. Mamm. i. p. 83 (1897: Batchian; Gilolo); 

Matschie, Mrr/ackir. pi. v. figs. 13. 14 (skull : Batchian) (1899). 
Pteropus affinis," Grm/, Cat. Monk. ^r. p. 108 (1870: Gilolo). 
Pteropus mackloti var. batchiana, Gray, Cat. Monk. ^-c. p 110 

(1870: Batchian). 
Pteropus (Acerodon^ muckloti b. batchianus, Matschie, 3Ipr/nchir. 

p. 10 (1899: Batchianl ; Trouessart, Cat. Mamm., Suppl. p. 49 

(1904: Batchian; Gilolo). 
Pteropus mackloti (pt., ncc Temm.), Dobson, Cat. Chir. B. M. p. 67, 

specimen a (1878: Batchian). 

Diagnosis. — Allied to Pt. dohsoni, but orbits larger, i., and p^ 
smaller, cheek-teeth slightly smaller, ears longer and more dir,- 
tiuctly attenuated above, general size sliglitly smaller, tibia much 
i.hortcr, and colour of head, mantle, and underparts much paler. 
Back and under.side of body dark brownish thickly varied with 
silvery buffy or buify straw-yellow, the ])ale hairs sometiraea nearly 
concealing dark colour on underparts ; head and neck ochraceous 
or ocbraceous-buffy : a mars-brown phase occurs. Forearm 135- 
1. 39-5 mm. Hah. Gilolo group; Sula Is. ; Sanghir Is. 

Skull.- — Similar in size and other characters to that oi Pt. dohsoni, 
but with conspicuously larger orbits ; orbital diameter 14*2-14"7 
mm., against 13 in Pt. dohsoni. — In size the skull of Pt. caniveps 
equals that of the largest races vi Pt. hypomelanus, from which it 
differs chiefly in the heavier, particularly broader, rostrum, broader 
palate, and larger orbits; ma\illary width across external sides 
of m^-m' 20-2-20-8 ram., against 16'5-19 in all races of Pi. hypo- 
melanus ; width of mesopterygoid fossa 8'2-8'7 mm., against 
7-8-2; width of palate between p^-p^ 12 mm., against 9-11-7; 
orbital diameter 14-2-14-7 mm., against 12-7-13-2. 

Teeth. — Not quite as heavy as in Pt. dohsoni, and with con- 
spicuously smaller i, and p,. Similar in structure to those of 
Pt. hypomelanus, but canines longer, cheek-teeth markedly broader 
(compare measurements, p. 199, with those of Pt. hypomelanus, 
pp. 118 and 131). — Ciugulum of upper and lower canines narrow. 
Posterior ledges of upper and lower premolars short and ill-defined, 
thougli a minute postero-external basal notch is occasionally detect- 
able in p' and p^. m'- larger than m^. i^ once and a half or nearly 
twice the size of i^. p, once and a half the size of i^, subequal 
to ni. 


Ears. — Longer than in Ft. lujpomelanus and dohsonl ; iipper third 
distinctly attenuated, owiii^- to flatly concave emarginatiou of outer 
margin ; tip narrowly rounded off. 

Interfemordl. — Scarcely developed in centre. 

Fur. — Short, silky ; closely adpressed on back. Approximate 
length, back 10-12, mantle li-5-l-i-5, belly 11-5-13-0 mm. Width 
of hairy space of back about 41-47 mm. 

Fur of back extending as a narrow line of short, and closely 
adpressed hairs along upperside of humerus and proximal fourth 
or third of forearm. Tibia naked above, except for a few scattered 
hairs on proximal third. 

Colour. — Two colour phases occur; in both, the head, mantle, 
sides of neck, and foreneck arc oehraceous or ochraceous-buff or 
cream-buff, generally with extreme base of hair dark brown ; in 
one phase the dark brown colour of back, rump, breast, and belly 
is strongly mixed with, sometimes more or less completely con- 
cealed or replaced by, glossy silvery huffy or buffy straw-yellow ; 
in the other phase the back, rump, breast, and belly are uniform 
dark mars-brown or russet mars-brown. 

(1) Grizzled phase: 5 yg. ad. skin, Batchian, type of species 
( — Back and rump glossy silvery buffy heavily mixed 
with seal-brown and blackish hairs. On front half of dorsum the 
dark and pale hairs are nearly equal in number, on posterior 
portion of dorsum and on rump the pale is largely in excess of the 
dark clement. — Breast, belly, and flanks pale silvery greyish with 
a faint tinge of cream-buff; base of fur slight!}' darker. — Mantle 
and sides of neck buffy, shading to cream-buff on occiput and to 
ochraceous-buif in shoulder region ; foreneck slightly darker than 
sides of neck and suffused with pale wood-brown. Hairs of" mantle 
and sides of neck seal-brown at base. — Crown and forehead similar 
to mantle, all hairs dark brown at base. Sides of head and throat 
mixed cream-bufly and dark brov/n. 

A second skin ( 2 ad. skin, teeth much worn, Batchian, 
is similar to the type in the general style of coloration, but differing 
in many details : — While the dark-coloured hairs of back and rump 
are seal-brown or dark vandyck-brown as in the type, the pale 
hairs are different in tinge, huffy straw-yellow, and rather more 
thinly spread, nowhere completely covering the dark fur. — Breast, 
belly, and flanks dark vandyck-brown, approaching seal-brown, 
thickly mixed with hairs similar in colour to pale hairs of back, 
making general aspect of underside of body much darker than in 
typo of species. — Mantle, occiput, and sides of neck darker than 
in type, between ochraceous-buff and oehraceous, shading almost 
to russet in shoulder region; extreme base of fur dark brown. 
Foreneck similar to sides of neck, but thickly mixed with dark 
brown hairs. — Crown somewhat darker than mantle, approaching 
clay, with dark brown bases to the hairs. Sides of head and throat 
similar to foreneck. 

The type of Ft. affinis Gray (jjuv. skin, Gilolo, 
is oil the wliole similar to the second example described above, but 



palc-coloured hairs of back, rump, and uuderparts more similar in 
tinge to those of the tj'pe of the species. 

(2) Mars-brown phase: d a<i- skin, teeth almost unworn, 
Batchian, type of Pt. macUoti var. latcJiiana, Gray ( — 
Back and rump uniform dark mars-brown without any appreciable 
admixture of pale hairs. Breast, belly, and flanks uniform russet 
mars-brown, with a few silvery white hairs. Head, mantle, sides 
of neck, and foreneck as in the grizzled phase. 

Sexual differentiation. — Canines conspicuously longer and heavier 
in males than in females. Measurements taken on one male and 
two females : upper canines, vertical extent from alveolar border 
11 mm. (male) and 9 (females), antero-posterior basal diameter 
4-8 (male) and 3*7-3-8 (females); lower canines, vertical extent 
9*5 (male) and approximately 6-5 -7-5 (females, tips slightly worn), 
antero-posterior basal diameter 3"8 (male) and 3 (females). 
' Measurements. On pp. 198, 199, 

: ' Specimens examined. Sixteen, in the collections of the Leyden 
(oleven, list under synonyms above'), Berlin (one), and British 
Museums, including the types of the species and of Pt. affinis and 
J't. macMoti var. hatcliiana, Graj'. 

Bange. Gilolo group : Morotai, Gilolo, Ternate, Batchian ; Sula 
Islands : Sula Bessi ; Sanghir Islands : Siao. 

^'ype in collection. 

Nomenclature and history in literature. — Chiefly two facts have 
influenced the history of this species in literature, viz., the unusually 
great individual variation in the colour of the fur, and the strong 
external resemblance of the species, even in size and colour of fur, 
to certain forms of the Acerodon macMoti group. The former fact 
explains that this species was described by Gray under three 
different names, the latter its confusion, by Gray and other authors, 
wath Acerodon mackhti. — The type of Pt. caniceps, Gray (1870), is 
a young adult female (practically full-grown, forearm 135 mm.) 
of the grizzled phase, collected in Batchian by Dr. Wallace. The 
type of P^ affinis, Gray (1870), is a j'oung, not nearly full-grown 
male (forearm 112 mm.) of the grizzled phase, collected in Gilolo 
by Wallace, and differing from foregoing only in its much smaller 
size (entirely due to immaturity) and rather darker colour, owing 
to a slight predominance of the darker over the paler hairs, espe- 
cially on the underside of the body. The type of Pt. macldoti 
var. batchiana, Gray (1870), is an adult male of the mars-brown 
phaSe, collected in Batchian (type locality of Pt. canicep>s) by 
Wallace, and externally not unlike the Timor specimen rightly 
referr(!d by Gray to Pt. macMoti. — By Dobson (1878, Z. s. c.) 
Pt. caniceps, though catalogued as a distinct species, was considered 
" probably a hybrid between Pt. hypomelamis and Pt. maclloti " ; 
Pt. affinis he rightly put down as a synonym of Pt. caniceps, but 
the type oi Pt. mackloti \a,r. batcJiiana he referred to Pt. mackloti 
(specimen «). — Matschie considered Pt. mackloti var. batchiana a 
local form ("Abart") of '^ Pier opus {Acerodon) macMoti," with 
which he united, with a query, Pt. caniceps, " Dobson " (really 


Gray), while the name Pt. affinis, Gray, appears to have escaped liis 
attention. The skull figured by Matschie (/. s. c.) under the name 
Pt. caniceps is that of a young t'eniale of tiiis species collected in 
Batcliian by Prof. E. v. Martens (Berlin Museum, no. ?A72). — 
Tiic name Pi. ht/pomelanns in Jeutink's 'Catalogue osteologiqiie ' 
and 'Catalogue systeinatiquc' (l. s. c.) covers three species, Pt. hijpo- 
melanus, cankrps, and palU<las. 

Jteinarks. — Pt. atniceps is readily distinguished from Pt. hypo- 
mdatins by its heavier roslrura, broader palate, larger orbits, 
heavier cheek-teeth, and much larger and more narrowly pointed 
ears. The colour of the fur, though very variable individually, 
never (so far as the available material goes) closely approaches that 
of any form oi Pt. liypomelanus. — From tbe species of the Acerodon 
machloti group, to which it bears much external resemblance, 
Pt. caniceps differs widely in the structure of the clicek-tecth 
(no trace of inner basal ledge). — Its differences from Pf. dohs<>id 
have been summed up in the diagnosis above. 

a. cJjuv. sk.; skull. Gilolo. Dr. A. E. Wallace ( 

(Type of PL affinii^. Gr;iT. > 
I). $ yg. ad. sk. ; Batcbian. Dr. A. E. Wallace (iU.l.lU.I. 

skull. [C]. 

( Ti/pe of species.) 

c. c? ad. sk. ; skull. Batchiaii. Dr. A. E.' Wallace 

(Type of Pt. ■inackloii var. hatchiaim. Gi;iv.) 

d. $ ad. sk. ; skull. Batcliiau {Br. A. Tomes Coll. 7.1 1.243. 

Pi. Wallace). 

27. Pteropus argentatiis, Gray. 
Pteropus chrysoproctus (pt.), Dobson, Cat. Chir. B. M. p. 48. 

Pteropus argentatu.'?, Gray, List Mamin. B. M. p. 194 (1843) 
(nomeu nudum) ; id., Voy. ^ Suljjhtir,'' i. p. .30 (1844 : ? Aniboina) ; 
Schinz, fSyst. Verz. Stiug. ii. Nachtr. p. 1.5 (184.5: Amboina) ; 
Wm/ner, Schreber'.? Siiuq., Siqjpl. \. p. .590 (1853-55: ? Amboimi) ; 
Fitzinger, SB. Aknd. 'Wim, Ix. Abth. i. p. 412 (1870: ? Am- 
boina) ; Gray, Cat. Monk. ^-c. p. 106 (1870 : ? Amboina). 

Pteropus chrysoproctus (pt., nee Tenim.), Dobsoii, Cat. Chir. B. M. 
p. 48, specimen a (1878: Moluccas): Trotiessart, Rev. ^- Ma(,. 
Zool. (3) vi. p. 205 (1879) ; Trottessart, Cat. Mamm. i. p. 80 
(1897) ; Matschie, Megachir. p. 14 (1899). 

Diagnosis. — Similar to Pt. canictps. but orbits smaller, cheek- 
teeth much smaller, fur longer. Upper side of body dark brown 
heavily sprinkled with longer yellowish hairs, under side russe'c 
mixed with longer huffy hairs ; mantle and head buffy or ochra- 
ceous-buffy, forcneck tawny. Probably rather larger than 
Pt. caniceps. Hah. Uncertain ('? Amboina). 

Slcull and teeth. — Type .skull defective and not full-grown ; 
orbits smaller, palate relatively narrower than in Pt. caniceps. — 
Check-teeth much smaller than in Pt. caniceps, but not differing 



in structure, except perhaps in the slightly stronger posterior 
ledge of p^, Cingulum of u])per and lower canines well defined, 
m'' larger than m^. i., twice the size of i,. p, once and a half the 
size of i.„ suhequal to m^. 

Ears. — Ptather long (about 31 mm. from notch to tip), reaching 
hinder corner of eye; distinctly attenuated above, tips rather 
narrowly pointed. 

Wings. — Membranes arising about 20 mm. apart from sides 
of back. 

Interfemoral . — Depth in centre about 3 mm. 

Fur. — Longer than in Pt. caniceps. Longest hairs, back about 
16, mantle and belly 20 mm. Least width of furred area of back 
40 mm. Distribution of fur as in Ft. caniceps. 

Colour (type, al.). — Back and rump seal-brown thickly mixed with 
buffy straw-yellow hairs, producing the impression of a dark brown 
fur heavily sprinkled with longer yellowish hairs. — Breast, belly, 
and flanks rich russet sprinkled with buffy hairs ; extreme base of 
fur seal-brown. Anal region seal-brown thickly mixed with 
huffy. — Mantle, occiput, and crown buffy, washed with ochraceous- 
buff in centre of mantle and on crown, and shading through 
ochraceous on sides of neck and head, to tawny on foreneck and 

; External measurement's of Pteropus dobsoni and caniceps. 


Pollex, total length, c. u 

,, metacarpal 

,, 1st phalanx 

2nd digit, metacarpal 

„ 1st phalanx 

,, 2ud-3rd phalanx, c. u 
3rd digit, metacarpal 

,, 1st phalanx 

„ 2nd phalanx 

4tli digit, metacarpal 

,, 1st phalanx 

5th digit, metacarpal 

„ 1st phalanx 

,, 2nd phalanx 

Interfemoral in centre, depth . 

Lower leg 

Foot, c. u 


Pi. dobsoni. i 

Pt. caniceps. 

6 yg- ad. 1 

3 ad.* 

























































50- r. 



* Type of species (young adult, but practically full-grown) ; type of 
Pt. macJdoti var. batchiana (adult); and no. (aged). 
t Estimate (skins). 



Mcnsurements of skulls a)id tectk (j/' I'teropus dobsoni, 
Cciniceps, and argentatus. 

PL. . 


skull, total leiigtli to gnathioii 

,, jialatioii to incisive i'oramiiia ... 
,, front of orbit to tip of nasals ... 
„ width of brain-case at zygomata . 

,, zygomatic width 

,, width in', externally 

,, lachrymal width j 

„ width across canines, externally. 

,, pnstorbital constriction 

,, interorbital consiriction ' 

,, width of niesopterygoid fossa ... 

,, between p^-p^, internally 

,, between cingula of canines 

„ orbital diameter 

Mandible, length ' 

„ coronoid height 1 

Upper teeth, cm- I 

Lower teeth, c-m,, 

Upper incisors, combined width 

p', length 

,, width 

p*, lengt h 

„ width 

ni', length 

,. width 

m=, length 

,, width 

p,, length 

„ width 

P3, length 

„ width 

Pp le"gtli 

,, width 

ni,, length 

„ width ' 

m^, length ' 

„ width 

Wj, length 

,1 width 














3 t 

Ft. caniceps. 
I Skulls: 2 ad.* 
I Teeth : 2 ad., 
1 2 imm.* 






























22 8 

24 2 









14-2 14-7 
53 8 


























n. argciitatns 
cJ imm. 
T^" pe. 





















* Skulls : type of Tt. mackloli rar. 

.......o. ,j,jc VI i I., ,/niinn/w ..... ha(chiaiia, and no. (skull 

measurements of type of species excluded, owing to slight immaturity of 
specimen). Teeth : the same specimens, and type of species, and type of 
J't. affinis. 

t Abnormal in 8hf.))e. 



throat. Base of fur of mantle, sides of neck, and head seal-brown ; 
fur of foreneck scarcely darker at base. 

Measurements of teeth on p. 199. Measurements of skull and 
external dimensions omitted, owing to immaturity of specimen ; 
the actual length of the forearm of the type is ]35"o mm. (in the 
adult Pt. caniceps 135-139-5); as the fronto-parietal sutures of 
the type skull are unobliterated, the fully adult Pt. argentatus is 
probably slightly larger than Pt. canicejis. 

Specimen examined. One, the type. 

Ranfje. Type presumed by Gray to have been obtained in 
Amboina (Voyage of the ' Sulph\ir '). In his Catalogue of 1870 
(p. 106) Gray gives as habitat of the species " Amboyna? Island 
of Gobie ; New Guinea." The British Museum does not possess 
any specimen of Pteropus from Gobie. 

Type in collection. 

liemarks. — A full description of this species cannot be given, the 
only specimen known being not full-grown and in a not quite 
satisfactory state of preservation. It is certain, however, that 
Dobson was mistaken in putting it down as a young Pt. cliryso- 
proctus. Judging from the characters of the teeth, the shape and 
size of the ears, the quality and distribution of the fur, and the 
general style of the colour, Pt. argentatus appears to be related to 
Pt. caniceps. 

n. J imm. al. ; "? Amboina " (Voy. of Sir E. Belcher 
bkiiU. the 'Sulphur'). [P.]. 

{Type of species.) 

D. The Pteiwpcs rufls geoup. 

Species. — Five species (six forms j: Pt.rufus{rufus &ud princeps), 
conwrensis, sei/chellensis, aldabrensis, and niger. 

Range. — The Malagasy region. This is the only group of the 
geuus distributed over the whole of the Malagasy region ; three 
other groups are represented by single species of very restricted 
range, viz., the melanopogon in Johanna Island, Comoros (Pt. living- 
sionei), the hypomelaniis in the Mascarenes (Pt. suhniger), and the 
lomhocensis group in Rodriguez (P<. rodricensis). 

General cliaracters (compare tig. 1:2, ]). 217, skull and dentition 
of Pt. niger). — Skull typical Pteropine. Dentition without 
special modifications ; cingulum of canines rather broad ; posterior 
ledges of premolars well developed but not particularly strong, 
ij and Pj not enlarged, m^ and m' not more reduced than usual. 
Ears long (much reduced in the single Mascarene species), at- 
tenuated at tip, subacutely pointed ; interfemoral distinct in centre, 
neither very deep (as ia Pt. vampijrus) nor quite obliterated. Colour 
in all except the Mascarene species characterized by the light 
mantle, crown, and underparts contrasting with dark back and 
muzzle. Males without glandular neck-tufts. Size generally rather 
large, sometimes below medium (forearm 124-171 mm.). 

Subdivisions of group. — The five species fall into two, rather 


sharply separated subsections, the one ranging over the whoha 
of the Malagasy region exclusive of the Mascarenes, the other 
confined to Mauritius and Reunion : — 

Malagasy type. — Pt. rufus (Madagascar), comoroisis, seychellensis, 
aldahrensis. Chief characters (in contradistinction to Mascarene 
type) : ears very similar in shape to those of Pt. giganteus, tibia 
naked or thinly haired above, general style of colour as in 
Pt. giganteus. It is noteworthy that the Malagasy species (rufus) 
is relatively sharply differentiated from the Comoro and Seychelles 
species (cumorensis, seychellensis), and these again from the Aldabra 
species (aldabrensis), so that, in spite of the geography of the 
islands, the Comoro species is closer to the Seychelles than to 
the Malagasy species, and the Aldabra form the most peculiar 
of the four. The Malagasy species {rufus) is the largest and 
largest-eared form ; colour pure giganteus style ; differentiated 
into two races, a northern and central (rufus), and a south-eastern 
(princeps). The Comoro and Seychelles species differ from P/!. ?-«/ms 
in rather smaller size and markedly smaller ears ; Pt. seychellensis 
is closely similar to Pt. comorensis, except iu the much stronger 
admixture of greyish white iu the colour of the back and rump. 
Pt. aldahrensis is chiefly characterized by its much smaller si/e and 
by having the colour of the back conspicuously lightened with 
broccoli-brown or wood-brown. 

Mcoscarene type. — One species only, Pt. niger (Mauritius, 
lleunion). Principal characters (as compared with Malagasy 
type) : ears extremely small, nearly concealed in the fur ; tibia 
thickly clothed above ; fur longer ; underparts dark-coloured. 
Pt. niger is unique in the genus in its style of colour : light sides 
of back contrasting with dark mantle and spinal tract. Owing to 
its small ears, hairy legs, and rather rich fur, this species has 
hitherto always been widely, by some authors generically or sub- 
generically, separated from Pt. rufus, and associated with the other 
hairy-legged Pteropi (dasyynallus, formosus, anetianus, pselaphon, 
poliocephalus, &c.) ; its skull and teeth are, however, in every 
respect, even to trivial details, similar to those of Pt. rufus and 

Affinities of group. — The Pt. nifus accords with the Pt. hypo- 
melanus group in all essential characters of skull and dentition, 
and, so far as its typical members are concerned, also in the quality 
aud distribution of the fur ; it differs chiefly in the shape of the 
ears and rather more distinct development of the interfemoral. 
In the shape of the ears and general style of colour the Malagasy 
Pt. rufus bears much resemblance to the Indian Pt. giganteus ; this 
(together with the neighbouring, though widely separated, habitats 
of these species) accounts for the fact that for many years they 
were kept together as one species under the name Pt. edwardsi. 
The rufus and vampyrus groups differ, however, so essentially in 
dentition that any relationship between the groups would 
seem to be excluded. 


28. Pteropus riifus, /?. Geoff. 

Pterojms cdwardsi (pt.), Dobson, Cat. Chir. B. M. p. 53. 

(Synonj'iiis under the subspecies.) 

Diagnosis. — Rostrum of skull lojig. Posterior ledges of p'\ p.,, 
and p^ distinctly marked otf frooi teeth. Ears long, exposed, 
attenuated at tip, subacutely pointed. Tibia naked or very thinly 
haired above. Mantle, crown, and underparts buffy, orange buffy, 
or yellowish buft'y, strongly contrasting with seal-brownish muzzle, 
back, and rump. Size above medium : forearm 158-5-170-5 mm. 
Bah. Madagascar. 

Skull. — Typical Pteropine. Deflection of brain-case moderate, 
alveolar line if projected backward passing through base of par- 
occipital processes and upper margin of occipital condyle, llostrum 
long, slightly compressed ; anterior margin of orbit above front, or 
some point of front half, of m'. Orbits rather large, diameter 
greater than width of rostrum across alveolar borders of p'-p' ; 
postorbital processes long, strong, in aged individuals often sepa- 
rated by a minute space from corresponding jjrocesscs on zygoma ; 
base of postorbital processes raised considerably above level of 
frontal plateau, making this latter between orbits deeply concave. 
Zygomatic arches flaring posteriorly ; sagittal crest strong or even 
very strong. Coronoid process rather high, but somewhat sloping ; 
coronoid height of mandible distinctly less than length of lower 
tooth-row, c-m.,. 

Teeth. — No special modifications. Upper canines slightly re- 
curved; cingulum broad, forming a sharply defined shelf at inner 
and posterior base of tooth, p' a minute terete spicule, early deci- 
duous. Posterior basal ledge of p^ short, postero-externally raised 
into a small tubercle separated by distinct notch from base of outer 
main cusp ; ledge of p' more or less obsolescent, though the postero- 
external notch is generally detectable; hinder border of p* vertical 
on longitudinal axis of tooth. m^ subcircular or subsquarish, 
usually slightly larger than m^ — i,, twice or twice and a half the 
bulk of ij. Canines recurved, cingulum well developed but narrower 
than in npper ones. Pj about twice the bulk of i.,. Posterior 
ledges of p^ and p^ long (longest in Pj), marked off postero- 
externally by a rather deep notch from base of outer main-cusp, and 
usually developing a more or less distinct postero-external tubercle; 
in m^ and m„ the ledge is shorter, but postero-external notch almost 
always easily detectable, m^ slightly larger than p,. 

FaJate-ridges. — 5-|-6-f3. First ridge terminating laterally at 
front of canine ; second at back of canine ; third at front of p'^ ; 
fourth at back of p'' ; fifth at front of p^ ; sixth at front of m^ : 
seventh at middle of m^ ; eighth at m^; ninth to eleventh behind 
m- ; twelfth to fourteenth situated at palation border. 

Ears. — Long, exposed, quite or almost reaching hinder corner of 
eye when laid forward. Inner margin flatly and almost evenly 
convex from base to tip ; outer margin flatly convex in lower 

rxERorus rttfus. 203 

two-thirds, deeply concave in upper tliird ; tip strongly attenuated, 
subacutely pointed. Naked on both surfaces, except at base pos- 
teriorly and anteriorly and along basal half of outer and -inner 

Wmr/s. — About 30 mm. apart at origin from sides of back. 

Intcrfemoral.—^'eW developed in centre (depth about 15 mm.), 
but generally completely covered by overhanging hairs of rump. 

Fur. — Somewhat adpressed on back and rump, semierect on 
nape of neck. Length moderate, 19-22 mm. on back, mantle, and 
belly. Width of furred area of back about 58 mm. 

Above, humerus and jiroximal third or half of forearm (except 
region round elbow, which is naked) covered with closely adpressed 
hairs. Fur of back extending on lateral membrane for about 
14 mm. beyond line of origin of membrane. Femur long-haired. 
Short, thinly spread hairs along outer side of tibia almost to ankle ; 
median upper surface of tibia naked on distal half, very thinly 
haired or almost naked on proximal half. Interfemoral long- 
haired in centre and laterally along proximal half of tibia ; inner 
margin and region above calcar naked. 

Below, antebrachial membrane, lateral membrane along outer 
side of forearm almost to carpus and between humerus and femur 
covered with rather long, woolly hair. Tibia and interfemoral (its 
central portion excepted) naked. 

Colour. — cJ ad. al., N. Madagascar, teeth almost unworn; : — Back and rump a shade of dark brown approaching 
seal-brown, thougli with a touch of vandyck-brown, chiefly owing 
to the very short paler brown (almost russet) tips to the hairs, 
these paler hair-tips rather more conspicuous on rump and inner 
side of tibia than on back. — Breast and belly yellowish-buff; 
individual hairs with long seal-brown bases and shorter yellowish- 
buff tips ; dark base of fur qnite or almost completely concealed on 
breast and belly, more or less showing through in anal region, 
giving this latter a conspicuous brownish tinge ; flanks dark brown 
varied with yellowish-buff tips to the hairs. — Mautle buff-yellow, 
shading into deep orange-buff on sides of neck and foreneck ; short 
concealed base of hair everywhere seal-brown. — Buff-yellow colour 
of mantle shading gradually into yellowish buff on occiput, crown, 
interocular space, and temporal region ; base of fur seal-brown. 
Muzzle, as far back as region immediately above and behind eyes, 
lower jaw, chin, and throat seal-brown, strongly contrasting with 
yellowish-buff crown. 

Individual variation in colour. — Not great, chiefly dependent on 
the greater or less intensity of the colours. Hack and rump some- 
times almost pure seal-brown (short brownish hair-tips completely 
absent), sometimes distinctly " powdered " with a paler brown 
(brownish hair-tips more conspicuous) ; an extremely thin sprink- 
ling with silvery whitish hairs occasionally detectal)le on close 
inspection. — Bright colour of breast, belly, and crown varying from 
pale glossy yellowish buff (almost buff) to a deep ochraceous-buft". 
approaching ochraceous.^Mantle varying from buff-yellow through 
ochraceous-buff to tawnv. 


The two subspecies described below do not differ appreciably in 

Haw/e. Madagascar. 

Ilahits. — Bats of this species are particularly common in the 
coast regions of Madagascar. They usually pass the day in the 
thickets of isolated islets near the coasts, suspended from the upper 
branches of tall trees ; observers agree that even in broad daylight 
a colony of Ft. rufus is difficult to approach, they generally become 
restless, many taking to flight with shrill cries, others climbing 
about from branch to branch, using the long claw on their -wings as 
-well as their hind feet. Early in the evening they may be seen 
passing at considerable heights, flock upon flock with short in- 
tervals, on their way from the sleeping-places to the feeding- 
grouuds. Their flight is straight, steady, somewhat crow-like, 
though rather heavier and with more frequent flappings of the 
wings. Skimming over water they often dip down to touch the 
surface. They feed chiefly on wild dates, on which they grow 
immensely fat. Large numbers are killed by the natives for food. 

Subspecies. — Two, differing in size only ; the one (rufus) dis- 
tributed over North and Central Madagascar, the other {princess) 
known from the extreme south of the island. 

Ke)j to the Subspecies of Pteropns rufus. 

a. Averaging smaller : skull, total length 69-74, 

forearm 158-5-lGo-5 mm. (N. and C. 

Madagascar) Pt. r. rtifiis, p. 204. 

h. Averaging larger : skull about 77, forearm 

170-5 mm. (3. Madagascar) Ft. ?: princeps, p. 208. 

28 a. Pteropus rufus rufus, E. Geoff. 

Fany, Flacowt, Hist, de Madagascar, p. 166 c. fig. (animal) (1658). 
Great Bat from Madagascar, Edwards, Nat. Hist. Birds, pt. iv. p. 180, 

pi. 180 (head, life size; animal, much reduced) (17-51). 
Vespertilio facie canina, Klein, Quadr. Disposifio, p. 62 (1751). 
Vespertilio vampyrus (pt.), Linn. Syst. Nat. 10 ed. i. p. 31 (1758) ; 

id., op. cit. 12 ed. i. p. 46 (1766) ; Boddaert, Flench. Anim. i. p. 68 

Pteropus vampyrus, HUger, Abh. Akad. Bei'liii, 1804-11, pp. 78, 84 

(1815: E. African Islands). 
Temate Bat (pt.), Pennant, Syn. Quadr. p. 359 (1781). 
Vespertilio caninus (pt.), Blumenbach, Hnndb. Naturq. 6 ed. p. 73 

Pteropus rufus, E. Geoffrey, Cat. Mamm. Mus. Nation. d'Hist. Nat. 

p. 47 (1803 : Madagascar). 
Pteropus, E. Geojfroy, Ann. Mus. d'Hist. Nat. xv. p. 92 

(1810: Madagascar); t'uvier, Reggie Anim. i. p. 123 (1817: 

Madagascar); Des7narest, Mamm. i. p. 109, no. 138 (1820: 

Madagascar); Schinz, Thierr. i. p. 154 (1821: Madagascar); 

Lesson, Man. Mamm. p. 109, no. 278(1827: Madagascar); 

Destnarest, Did. Sci. Nat. xlvi. p. 359 (1827 : Madagascar) ; 

Gi-ay, hi Griffith's Anim. Kingd. v. p. 55, no. 154 (1827 : Mada- 
i gascar) ; Is. Gcofroi/, Diet. Class. d'Hist. Nat. xiv. p. 699 (pt.) 


(1828: Madagascar); Temmmck, Moii. Maimn. ii. p. 61 (pt.) 
(18o7: Madaiiasoar) ; Wagner, Schref/er'n Siiuf/., Si/ppl. i. p. 345 
(pt.) (1839 : Madflgascar) : Lessoti, N. Tabl. Ii. Anini., Mamm. 
p. 12, no. 169 (pt.) ('1842: Madagascar); Schinz, Syst: Verz. 
Sdiiff. i, p. 121 (])t.) (1844: Madagascar); IS. Desmarest, Diet. 
Univ. d'llid. Nat. xi. p. 247* (pt.) (1848: Madagascar); 
Grat/, Zvol. ' '^atnaratiff,' Vert. p. 11 (pt.) (1849: Madagascar); 
Wagner, Schreber's Sdiig., Stqypl. v. p. 595 (pt.) (1853-55 : 
Madagascar) ; Giebel, Sdug. p. 995 (pt.) (1855 : Madagascar) ; 
A. Newton, P. Z. S. 1865, p. 833 (Moliambo) ; Verreaux, Viusoyis 
Voy. u Madcu/ascar, Annexe A, p. 1 * (18(35) ; Schlegel, P. Z. S. 

1866, p. 419" (pt.) (Madagascar) ; Peters, M.B. Akad. Berlin, 

1867, p. 325 (pt.) (Madagascar); Pollen, in Pollen 8)- v. Dam, 
Peek. Fawie Madagascar, ii. pp.25, 164, 167,172 (pt.)(1868: 
Madagascar; habits); Fitzimjer, SB. Akad. Wien, Ix. Abth. i. 
p. 416 (pt.) (1870: Madagascar); Grainge, Anfa?ia7iarivo Ann. 
i. pt. 1, p. 25 (1875: Majunga, N.W. Madagascar; habits); 
Dobson, Cat. Chir. B.M. pp. 53, 55 (pt.) (1878: Vohemar, N.E. 
Madagascar) ; Trouessart, Rev. Sc Mag. Zool. (3) vi. p. 202 (pt.) 
(1879: Madagascar); Dobson, Rep. Brit. Assoc. 1878, pp. 162, 
165 (1879: remarks on distr.) ; Trouessart, Ann. Sci. Nat. (6) 
Zool. viii. Art. 12, p. 15 (1879 : remarks ou distr.) ; Gill, Stand. 
Aat. Hist. V. fig. opposite p. 102 (1884) ; Oliver, 3Iadagascar, i. 
p. 520 (1866); Jentink, Cat. Ost. Mamm. p. 259 (pt.) (1887: 
Madagascar) ; id., Cat. Sy.^t. Mamm. p. 146 (pt.) (1888 : Mada- 
gascar) ; Trouessart, Cat. Mamm. i. p. 81 (pt.) (1897 : Mada- 
gascar) ; Sibree, Antananarivo Ann. vi. pt. 21. pp. 33, 47 (1897 : 
Madagascar ; liabits) ; Lorenz-Liburnau, Abh. Senck. nat. Ges. xxi. 
pt. 3, p. 455, pi. xxxii. ligs. 1 «, 1 6, 2 (skull) (1898: Amburvi, 
N. of Majunga); Keller, Qstafr. Inseln, pp. S2, 121 (1898: 
Madagascar; Nossi Be) ; Matschie, Megachir. p. 15 (pt.) (1899: 
Madagascar) ; Grandidier, in Blanchard, Madagascar, p. 172, 
fig. 84 (animal) (1902) ; Trouessart, Cat. Mamm., Stippl. p. 50 
(pt.) (1904: Madagascar); Miller, Fam. S,- Gen. Bats, p. 58 
(1907) ; Grandidier, Noms Malgaches d'animait.v, p. 9, c. fig. p. 15 
(animal) (1908). 

Pteropus madagascariensis, Oketi, Lehrh. Natvrg. iii. Abth. ii. p. 936 
(1816: Madagascar). 

Pteropus phaiops, Temminck, Mon. Mamm. i. p. 178 (1825: Mada- 
ga.'5car) ; Lesson, Man. Mamm. p. 110, no. 285 (1827 : Madagascar); 
Desmarest, Diet. Sci. Nat. xlvi. p. 360 (1827: Madagascar); Is. 
Geoffroy, Did. Class. d'Hist. Nat. xiv. p. 700 (1828 : Madagas- 
car) ; Anonymous [? Vigors ^- Gould\ Cat. 3Ia7nm. Mus. Zool. 
Soc. p. 10, no. 162 (1828: Madagascar); id., Cat. Anim. Mus. 
Zool. So'.: p. 11, no. 162 (1829: Madagascar): J. B. Fischer, 
Syn. Mamm. p. 82, no. 4 (1829: Madagascar); Lesson, Hist. 
Nat. Mamm. (Compl. Buffon) v. p. 52 (1836: Madagascar); 
Temminck, Mon. Mamm. ii. p. 66 (pt.) (1837 : "Macassar," 
really Madagascar) ; Waterhouse, Cat. Mamm. Mus. Zool. Soc. 
J). 13, no. 100 (1838: Madagascar); Gray, Mag. Zool. 8,- Bot. 
li. p. 502 (1838 : Madagascar) ; S. Miiller, in Temminck, 
Nat. Ge.wk. Nederl. Overz. Bez.. Zoogd. pp. 20, 59 (pt.) 
(1839-44: "Celebes," really Madagascar); Lesson, N. Tabl. 
R. Anim., Mamm. p. 13, no. 171 (pt.) (1842: "Macassar,"" 
really Madagascar) ; E. Desmarest, Diet. Univ. dllist. Nat. xi. 

* )lis!!pelt r/eroptts edwar.tii. 

206 PTEUorus eufus eukus. 

Gervais, Hift. 

^ , „^. u, i.^^. ^*.,.,^. ^~. , ^w„. , J,. — , V -~ — -„..., .. 

Madagascar^ ; Matschie, Megachir. pi. i. tigs. 4, 4 a, 4 6 (skull) 

La C-irande Eoussette (Fsnii), Sr/anzin, Mem. 8oc. Mvs. d'Hist. Nat. 

Strassboui-f/, iii. pt. i. Mt^m. MM. p. 11 (1840: habits"). 
Grosse chauve-soaiis noire et jaiine, Laillet, Madagascar, p. 29 


Diagnosis. — Skull arid external dimensions smaller (see measure- 
ments on pp. 210, 211). Forearm 158-5-165-5 mm. //rtS. N. and 
Central Madagascar. 

Specimens examined. — Eight, in the collections of the Lej'den and 
British Museums, including the cotypes oi Pt. phaiops, Temm. 

Banr/e. — North and Central Madagascar : Majunga, Amburvi, 
Nossi Be, Vohemar, Fianarautsoa. 
Ty2:>e probably not in existence. 

Earliest histori/ in literature. — The records of this species in 
literature date back at least to the first French attempts at a 
colonisation of Madagascar, about the middle of the seventeenth 
century; thus ib was briefly described and figured by Flacourt 
(1658, op. s.c, chapter " Oyseaux de nuict") under its native name 
" Fany " (by later writers usually spelt Fani, Fanny, Fanii, or 
Fanihy) : Flacourt's illustration is probably the most imaginative 
figure ever given of a bat. A tolerably good coloured figure of the 
head of this species was published, nearly a century later, by 
George Edwards in his 'Natural History of Birds' (1751, I. s. c). 
It is one of the three species covered by the Linnean name 
Vespertilio vampyrus (1758 and 1766, sec infra, pp. 219, 351). 

Pterojms rufus, E. Geoff. ; 180^. — Type locality, Madagascar. 
Based on an " individu envoye par le citoyen Mace, naturaliste " 
(no. 90 of Geoffroy's Catalogue). I have been iinable to find this 
specimen in the Paris Museum, but Geoffroj's brief description of 
its colour leaves no doubt whatever as to the identification of the 
species. Only reference given by Geoffrey : Edwards, Great Bat 
fi'om Madagascar. 

Fieropus eclwarclsi, E. Geoff. ; 1810. — Type locality, Madagascar. 
In reality a renaming and redescription of the same author's 
Pt. rufus, based ou an example " dont nous sommes redevables a 
I'estimable naturaliste M. Mace," therefore most probably a re- 
description of the very type of Pt. rufus (see above). Only 
references: Edwards and Linnasus (F. vampyrus). — Temminck, in 
1825 (Mon. Mamm. i.), put Pt. exlwardsi down as a synonym of 
Pt. edidis (i. e. Pt. vampyrus), while at the same time he described 
the true Pt. edwarchi under the name Pt. phaiops, and Pt. yiganteus 
(Continental India) as Pt. meclius ; in 1837 (Mon. Mamm. ii.) he 
recognized the validity of the Geofl'royan Pt. edwardsi, put his own 
Pt. medins down as a synonym of this species, which therefore he 

* yi\sB]>e]t P/i'roft/s JlJia?ir:pS. 

PTERorus R0Frs RVFrs. 207 

considered " repauduc dans toute Flnde, a Ceylon et a Madagascar," 
and deliberately cliani;-ed the type locality of Ft. I'luiiops from 
" Madagascar "' into " Macassar," erroneously believing the former 
word to be a slip of the pen for the latter. It is of some importance 
to bear this confusion of names and species in remembrance, 
because Temminck's view as to the specific identity of Pt. edwardsi 
and the ratlier similarly coloured Pt. meditts was accepted by a 
long series of later writers (see synonymy of Pt. (jiganteiis), so that, 
when no locality is given ( e. g. in anatomical papers), it is often 
difficult or impossible to see if the name " Pt. edwardsi " refers to 
the Malagasy {Pt. nifus) or the Indian species {Pt. giganteus). 

Pteropus madagascarlensis, Oken • 1816. — Type locality, Mada- 
gascar. A renaming of E, Geoflfroy's Pt. edicardsi. Eefercnces : 
Pt. edwardsi and Vespertilio vampyms. Description an almost 
verbal German translation of Geoffrey's description (size, colour) of 
Pt. edwardsi. 

Pteropus pliaiops, Temm. ; 1825. — Temminck's Pt. phaiops, 
1825, is E. Geoffroy's Pt. rufus; Temminck's Pt. jjJiaiojjs, 1837, is 
a mixture of Pt. rufus and Pt. melanopogon. The explanation of 
the mistake is this : In 1825 Temminck described two specimens 
of a Pteropus from Madagascar as Pt. phaiops: both of these 
specimens are still in the collection of the Leyden Museum ( S ad., 
$ ad., mounted; Jentink, Cat. Syst. p. 146, sub Pt. edwardsi, 
specimens c, d; skull of c in situ, skull of d separate: Cat. Ost. 
p. 259, c) ; they are in every respect indistinguishable from 
British Museum specimens of Pt. rufus rufus ; but when later on 
receiving six s])eciraens of a Pteropus from Amboina, obtained by 
Mliller and Macklot, Temminck believed in these to recognize his 
Pt.pJutiops, and considering that the same species could hardly be 
common to Madagascar and Amboina, he now (Mon. Mamm. ii. 
p. 66, 18.37) regarded "Madagascar" as an " erreur ou faute 
d'impression " for " Macassar " (Celebes). The truth is that the 
cotypes of Pt. pJiniops were correctly ticketed Madagascar, while 
Miiller and ilacklot's Amboina specimens were the very different 
Pt. melanopogon; that Temminck failed to distinguish the latter 
from the former is easily explained by the fact that Pt. meJano- 
pogon in the colour of the fur bears no small resemblance to 
Pt. rufus. — All later records iu literature of Pt. phaiops, rcsj). 
phcfops, from ^lacassar or Celebes in general (see references above) 
are based solely on Temminck's deliberate changing of the true 
type locality of Pt. pJiaiopis, viz. Madagascar, into Macassar. 

a. A. 2 ad. sks. ; Jfadagasfar. Purcli.ised (Blyth,,29. 

skulls. Greene & Jourtlain I. 

f. (Jad. al. N. M:iclag:iscar. Rev. R. Baion [C. & 

(/. Ad. skeleton. N.Madagascar. A. Crossley [C.]. 70.5..5.1I. 

(. Ad. sk. ; skull. Vohemar, K.E. A. Crossley [C.]. 70..').5.4o. 

/. 9 iiTi"'. liead : Finnarantsna, Eoyal Society [P.]. 97.91. 29. 
pkiill. Central Mada- 

gascar ( Dr. C. f. 
I'uis-iilli Major). 


28 h. Pteropus rufus princeps, K. And. 

? Pteropus edwardsi {nee K Geof. subsp.), Burtlett, P. Z. S. 1875, 
p. 63 (S.E. coast of Madaoascar). 
Pteropus rufus princeps, K. Andeisen, Ann. S,- Mag. W. H. (8) 
ii. p. 367 (] Oct. 1908: Ft. Dauphin). 

Diagnosis. — Skull and external dimensions larger (see measure- 
ments, pp. 210, 211). Forearm about 170'5 mm. Bob. S. 

Specimen examined. One, the type. 

Ranr/e. South Madagascar : Ft. Dauphin. 

Type in collection. 

o. c^ ad. al. ; skull. Ft. Dauphin, S.E. M. Cloisel [C.]. 
Madagascar. {lyv^ of subspecies.) 

29. Pteropus comorensis, Nieoll. 

PteropitrS edwardsi (pt.), Dobson, Cat. Chir, B.M. p. 53. 

Pteropus edwardsi (nee E. Geoff.), Peters, Seise Mossambique, Zool. 
i. Siiug. p. 23 (1852: Johanna 1.; notes on anatomy ; habits); 
Wagner, Schreber^s Siiuq., S'lippl. v. p. 595 (pt.) (1853-55: 
Comoros) ; P. Z. Sclaler, P. Z. S. 1861, p. 268 (Johanna) ; 
Schlegel, P. Z. S. 1866, p. 419 (Mayotte) ; Peters, 3IB. Akad. 
Berlin, 1867, p. 325 (pt.) (Comoros) ; Pollen, in Pollen 8^ v. Dam, 
Rech. Favne Madagascar, ii. pp. 25, 164 (1868; Mayotte; 
habits) ; I'Xtzinger, SB. Akad. Wien, Ix. Abth. i. p. 416 (pt.) 
(1870: Comoros); Gray, Cat. Monk. S,c. p. 103(1870: Shoa, 
errore ; Mohilla; Johanna); Dobson, Cat. Chir. 5. il/. pp. 53, 55 
(1878: Mohilla; Johanna) ; Troiiessart, Rev. (§• Mag. Zool. (3) 
vi. p. 202 (pt.) (1879: Mohilla; Johanna); Dobson,' Rep. Brit. 
Assoc. 1878, p. 162 (pt.) (1879: remarks on distr.) ; Jentink, 
Cat. Ost. Mam?)!, p. 2G0 (1887 : Mayotte) ; id.. Cat. Sgsf. 
Mamm. pp. 246, 247 (1888: Mavotte) ; Milne-Rdwards ^ 
Oustaht, N. Arch. Mns. d'Hist. Nat' Paris, (2) x. p. 223 (1888 : 
Mayotte ; Mohilla ; Johanna) ; Brehm, Tierleben, 3 ed. i. p. 344 
(1890 : Mayotte ; habit.^) ; Tronessart, Cat. Ma7mn. i. p. 81 (pt.) 
(1897: Mohilla; Johanna); Matschie, Megachir. p. 15 (pt.) 
(1899: Angazija ; Johanna); Trouessart, Cat. Mamm., Suppl. 
p. 50 (pt.) (1904 : Comoros). 
Pteropus rubricollis [nee E. Geoff.), Layard, Cat. S. Afr. Mus. p. 19 
(1861: Johanna). 
^ Pteropus comorensis, Wallace, Island Life, p. 400 (1880: Comoros) 
(nomeu nudum) ; id., op. cit. 2 ed. p. 428 (1892 : Comoros) (nom. 
nud.) ; Keller, Ostafr. Inseln, p. 125 (1898: Comoros) (nom. 
nud.) ; Nieoll, Three Voy. of a Naturalist, pp. 87, 88, 90 (1908 : 
Mayotte; habits). 

Diagnosis. — Similar to Pt. rufus, but smaller, and -with relatively 
much smaller ears and shorter hind legs. Forearm 151-157 mm. 
Hah. Comoro Is. 

Slcidl and teeth. — Differing only in smaller size from those of 
Pt. rafus rufus. Ten skulls : total length 65-5-68 mm. (69-73-8 in 
five skulls of rafi(.'<); mandible 50-8-53-8 (54-5-58-2) ; diameter of 


orbit 12-8-13-2 (13-8 14-6). Maxillary tooth-row, c-m% in ten 
specimens 24-25'0 mm. (26--27'8 in five rtiftis) ; m' in 18 specimens, 
length 5-5-5 (5-7-5-8 in six rufus), breadth 3-3-1 (3-1-3-6); m„ 
length 4-5-4-9 (5-1-5-4), breadth 2-8-3-1 (3-1-3-2). For further 
details sec measurements on p. 211. 

Ears. — Much smaller than in Pt. rufu/s, but not differing in 
shape ; length from lower margin of orifice about 31-5 mm. (37-38 
in rufus). 

Fur. — Distribution, qualitj', and length of fur as in Pi. rufus. 

Colour. — Three cJ ad. skins, teeth almost unworn, Mayotte, co- 
types of species, : — Back and rump blackish or blackish 
seal-brown, very thinly sprinkled with silvery greyish-white hairs. 
Head, mantle, and underparts as in Pt. rufus (p. 203). 

Specimens from Johanna, Moliilla, and Angazija do not differ 
appreciablj- in colour. The blackish or blackish seal-brown colour 
of the back fades almost invariably in the course of a few years to 
vaudyck-brown, even in specimens not exposed to light. 

E.vlernal dimensions. — Smaller, at least on average, than 
Pt. rufus rufus, with proportionally much shorter tibitc. Forearm 
of twelve adults 151-157 mm. (158-5-165-5 in six adult ru/ws) ; 
third metacarpal 99-105-5 (109-113); lower leg 68-72 (80-82); 
foot with claws 47-50 (53-57). For details see measurements on 
p. 210. 

Sj^ecimetis examined. Nineteen, in the collection of the British 
Museum, including the cotypes of the species, and representing all 
ihe islands from which the species has been recorded. 

Ranr/e. Comoro Islands : Mayotte, Johanna (Anjuan), Mohilla, 
Angazija (Great Comoro). 

Cotijpes in collection. 

Pteropus comorensis, Nicoll ; 1908. — The name Pt. comorensis 
seems to have been introduced into literature by Wallace in his 
' Island Life' (1880, I. s. c.) ; at least I have been unable to trace 
the name further back ; neither in the first nor in the second 
edition of that book (1892) does Wallace give any description ; he 
appears to be unaware that he was using a new name for the 
Comoro bat. Keller (1898, I. s. c.) probably copied the name from 
Wallace. Nicoll admittedly (in litt. to the present writer) found 
the name in Wallace's book atid, inadvertently, made it technically 
valid by the following descriptive note : " its fur was of a reddish 
colour, thick and soft " (1908, I. s. c. p. 88). 

«. <5 ad. sk ; Mayotte, Oomoros, sea- Earl of Crawford ft.6.3.1(>. 
Bkull. level; 25 Feb. 1906 [P.]. 

(M. J. Nicoll). 
i,/-. 2cJad. Bks.; Mayotte, 600' ; 2 Mar. Earl of Crawford,15. 
skulls. 1906 (M. J. Nicoll). [P.]. 

(a-c, cotypes of species.) 
d. ^ ad. al. Mayotte (M. J. Nicoll). Earl of Crawford 

*. Imm. sk. ; Johanna I., Comoros. John Barrow, Esq. 53,5.27.1. 
skuU. [P.]. 



Bewsher [C.]. 

. /. Yg. ad. sk.; Johanna !.(/);•. Zu>/»^- EarlEussell [P.]. 

skull. s/oiie}. 

ff- HI. 4c?ad.,l iS Johanna I. 
inim., 1 ad., 
1 imm. sks. ; 
n. 1mm. sk. ; Moliilla, Comoros ; 

skull. Aug. 1862. 

o. Imm. skull. Mohilla. 
])• Imm. sk. ; Angazija, Comoros. 
skull, skeleton. 
g-s. Imm., (j"yg. Angazija. 
ad, 2 ad. sks.; 

Sir J. Kirk [C.]. 

Sir J. Kirk rC.]. 
Sir J. Kirk [C.]. 

Sir J. Kirk [C.]. 

6.- bis. 

External measurements of Pteropus rufus and comoreiisis. 

Pt. rufus 

rt. cmnoreiisis. 


6 ad. 

(Incl. cotj'pes of 

Pt. 2}fiaiops.) 


12 ad. 
(Incl. cotypes.) 


MiN. Max. 

mm. mm. 
168-5 165-5 
68 72-5 
15-5 17 
35-5 37 
80 85 
19-5 21-5 
18 21 

109 113 
82 86 

111 118-5 

108 110 

64 70 

63-5 71 

110 116 
48 53 
51-5 66-5 




15 5* 
80 82 
63 57 
(19 1) 22-5 

















MiN. Max. 

mm. mm. 

151 157 
61-5 66-5 
14 10 
31 36 
72 78 

18 21 
16-5 18-5 
99 105-5 
76 81 

104 ] 16 
98 103-5 
61 65-5 
60 64 
101 107 
45 48 
47 54 




68 72 
47 50 

19 (20t) 

Pollex, total length, c. u 

„ metacarpal 

„ 1st phalanx 

2nd digit, metacarpal 

., 1st phalanx 

2nd-3rd phalanx, c. u. 
3rd digit, metacarpal 

„ 1st phalanx 

,, 2nd phalanx 

4tb digit., metacarpal 

,, 1st phalanx 

,, 2nd phalanx 

,, 1st phalanx 

,, 2nd phalanx 

„ greatest width, flattened ... 
Front of eje to tip of muzzle 
Interfemoral in centre, depth ... 

Ff)ot c. u 

* From one alcoholic speciiuen. 
■|' From dried skin. 



Measurements of skulls and teeth of Pteropus rufus and comoreusis. 

Ft. rufus 

Ft. comwensis. 

Skulls : 5 ad. 
Teeth : 5 ad., 

1 imm. 

(Incl. cotype of 

Ft. pluuops.) 


Skull: 10 ad. 
Teeth: Had., 
7 iram. 
(Incl. cotypes.) 

Skull, total length to gnathion 












































i Mix. 



. 1-8 




































,, piiliitiou to incisive foramina ... 
,, frout of oi-bit to tip of nasals ... 
„ width of brain-case at zygomata. 
„ zygomatic width 

„ width across m', externally 

„ lachrymal width ". 

„ widtii across canines, externally. 
,, postorbital constrit'tion 

„ interorbital constriction 

„ width of niesoptervgoid fo.ssa... 

„ between p'-p^, internally 

„ between cingula of canines 

., orbital diameter 

Mandible, length 

„ coronoid height 

Upper teeth, c-m- 

Lower teeth, c-m^ 

Upper incisors, combined width 

p^ length 

„ width 

p*, length 

„ width 

m', length 

,, width 

m', length 

„ width 

Pp length 

„ width 

p,, length 

„ width 

p,, length 

„ width 

ni,, lengtli 

„ width 

m„ length 

I, width 

tn,. length 

„ width 



30. Pteropus seycliellensis, Mllne-Edw. 
Pteropus edtuardsi (pt.), I*obson, Cat. Chir. B.M. p. 53. 

Pteropus edwardsi {n£c E. Geoff.), Wright, Ann. ^ Mag. N.H. (4) ii. 

p. 4-j6 (1868 : Seychelles ; habits) ; Wallace, Geogr. Distr. Anim. 

i. p. 281 (1876 : Seychelles) ; Dobson, op. s. c. p. 53 (pt.) (1878 : 

Seychelles); Trouessart, Rev. i^- Mag. Zool. (3) vi. p. 202 (pt.) 

(1879: Seychelles). 
Pteropus seycliellensis, A. Mibie-Edicards, Ball. Soc. Fkilom. (7) 

ii. p. 221 (1887: Seychelles); Trouessart, Cat. Mnmm. i. p. 81 

(1897: Seychelles) ; Miller, Fam. Sr Gen. Bats, p. 58 (1907). 
Pteropus edwardsi a. seychellensis, Matschie, Megachir. p. 16 (1899 : 

Seychelles) ; Trouessart, Cat. Mavnn., Suppl. p. 50 (1904 : 


Diagnosis. — Similar to Pt. comorensls, but dark colour of back 
and rump conspicuously sprinkled with silvery greyish-white. 
Hah. Seychelles. 

Skull and external characters, except colour of fur, as in 
Pt. comorensis ; teeth on the whole slightly smaller. 

Colour (four specimens, slightly immature ; cotypes). — Back and 
rump blackish seal-brown conspicuously sprinkled with glossy 
silvery greyish-white hairs, particularly on sides of back along 
membranes and on rump ; owing to the mixture of seal-brown and 
greyish-white the general impression of the colour of the rump and 
hinder back approaches hair-brown or even very light mouse-grey ; 
the degree of sprinkling with light-coloured hairs, though always 
conspicuous, is not equally strong in all specimens. — Breast and belly 
with long seal-brown bases to tho hairs and short ochraceous or 
ochraceous-buff tips ; these latter being too short to completely 
cover the dark bases, the general aspect of the colour is a dark 
brownish, lightened or thickly blotched with ochraceous or ochra- 
ceous-buff ; some long shiny silvery greyish- white hairs on breast 
and belly. Flanks and anal region more uniform blackish seal- 
brown, some of the hairs with short ochraceous tips ; a slight 
sprinkling with greyish-white hairs. — Mantle, occiput, crown, and 
sides of head and neck golden ochraceous-buff, in many places 
deepening to rich golden ochraceous, all the hairs with concealed 
blackish seal-brown bases ; foreneck similar, or clouded with pale 
russet. Muzzle, chin, and throat blackish, as in Pt. rufus and 

As will be noticed from the above description, the only essential 
difference in colour from Pt. rufus and comorensis is the pronounced 
admixture of greyish-white on back and rump, and the darker 
colour of breast and belly, this latter owing to the bright tips of the 
hairs being shorter, and the dark ground-colour of the fur therefore 
more exposed. — 

Measurements. On pp. 221, 222. 

Specimens examined. Eight, in the collections of the Paris (five), 
Berlin (one), and British Museums (two), including the cotypes of 
the species. 


liange. Seychelles. 

Coty^ies in the Paris Museum. 

Habits.— Ft. seychelleiisis is very common on all the islands of 
the Seychelle group. Though often on the wing in strong sunlight, 
between 8 and 10 in the morning, it is chiefly nocturnal in its 
habits. About an hour before sunset these fruit-bats may be seen 
flying at great heights from their resting-place in the woods, making 
as if they were going to cross the island and then, Avhen just over 
the group of trees to be visited, falling down as it were among 
them. The first comers take up good places, with plenty of fruit 
near them, alighting without noise ; by-and-by the arrivals are 
more numerous, and then the noise begiiis, every late comer trying 
to dislodge an earlier comer, and this not without much growling 
and grumbling and chattering. A little after sunset the noise is 
generally at its highest, and more than a hundred bats may now be 
found feeding in one small group of trees. Their favourite food 
seems to be the " fruit de Cythere " {Spondias cytherea) or the 
Mango {^Manyifcra inclica), but almost any fruit is welcome to them. 
The flesh of these Bats is much appreciated by the Creole inhabitants 
of the Seychelles. (Ed. Perceval Wright, I. s. c.) 

Pteropus seychellensis, A. Milne-Edw. ; 1378. — Type locality, 
Mahe, Seychelle Islands (Lantz coll.). The Paris Museum possesses 
a series of specimens obtained b)' Lantz in Mahe, at least four of 
which must be regarded as cotypes of the species, viz. three mounted 
specimens (group on one block), all slightly immature, and one 
2 iram., unmounted skin ; skulls of all in situ ; llcg. nos. 1106, 
1108, 1109, 1112. Separated by Milne-Edwards from " Pi. ed- 
u'ctrdsi " from Madagascar on account of its darker-coloured under- 
parts, and because " le collier roux ferrugineux si marque de ccs 
chiropteres [ i. e. edivardsi] manque chez ceux des Sej'chellcs." I am 
unable to explain how Milne-Edwards came to give this latter 
character; as a matter of fact, all the specimens from the Seychelle 
Islands, including those examined by Milne-Edwards, are similar 
in the colour of the mantle to Pt. rufus and comorensis. 

a. S P"ll. al. Seychelles. Dr. Ed. Perceval Wright [C.]. 1. 

A. 5 imni, al. ; skull. Seychelles. J. S. Gardiner, Esq. [P.]. 

31. Pteropus aldabrensis, True. 

Pteropus aldabrensis, True, Proc. U.S. Nat. Mus. xvi. p. 533 

(U Julv, 1893: Aldabra I.); Abbott, torn. cit. p. 762 (1893: 

Aldabra") ; Trouessart, Cat. Manun. i. p. 81 (1897 : Aldabra) ; 

Lorenz-Liburnaii, Abh. Senck. nat. Ges. xxi. pt. 3. p. 455, pl.x.xxii. 

fig.^. 3 a,Sb,3c,3 d (sku\\) (1898 : .\ldabra) ; VoeltzI;ow, op. cit. 

XX vi. pt. 0, pp. 543, 552 (1902 : Aldabra) ; Mdler, Fam. S,- Gin. 

Bats, p. 58 (1907). 
Pteropus edwardsi h. aldabrensis, Matschie, Megachir. p. 16 

(1899) ; Trouefsart, Cat. Mamvt., Suppl. p. 50 (1904: Aldabra). 
Plcropus [sp.1. VoeltzUou; Abh. Senck. nat. Ges. xxi. pt. 1, p. 66 

(1897 : Aldabra). 

Diagnosis. — Allied to Ft. coniorennis, but much smaller, and with 


tlio colour of the back and rump strongly tinged witli broccoli- 
brown and wood-brown. Forearm 134-5 mm. Ilab. Aldabra I. 

HhiU. — Not differing from that of Pt. comorensis except in the 
much smaller size and relativelj' much broader mesopterj'goid 
fossa ; Avidth of this latter 7"6 mm. {i. e. nearly seven-eighths of 
palate Avidth between fronts of p*-p^), against 7'2-7'8 in the much 
larger Comoro species (i. e. seven-tenths of palate width between 
fronts of p^-p''). 

Teeth. — Structure and relative size of teeth quite as in Pi. co- 
incrensis and allied species, except perhaps for a very slight reduction 
of m'^ and m^. 

Ears. — Shape and relative size as in Pt. comorensis. 
Fur.- — Eather softer, more silky than in Pt. comorensis. Approx- 
imate length of longest hairs of back, mantle, and belly 21 mm. 
Width of furred space at middle of back (measured on dried skin) 
64 mm. Distribution of fur as in the Comoro species. 

Colour. — cJ ad., teeth slightly worn, skin: — General colour of 
back and rump a shade of broccoli-brown, darkest on front and 
middle of back, palest (almost wood-brown) on rump. Majority of 
individual hairs of back pale greyish at base vfith wood-brown 
extremities. A rather conspicuous sprinkling with silver}' greyish- 
white hairs, particularly on sides of back, along membranes. Hairs 
on forearm and sides of tibia mars-brown. — Breast and belly ochra- 
ceous-buff with a tinge of bufi-yellow ; long concealed bases of 
hairs seal-brown. Flanks more uniform dark broAvn owing to 
shortness or, in many places, complete absence of bright-coloured 
tips. — Mantle orange-buff strongly tinged with ochraceous-rufous, 
shading posteriorly, in a transverse line across shoulders, into bright 
ochraceous-bufF, and on sides of neck and foreneck into deep 
ochraceous-rufoua. Concealed base of hairs of mantle and foreneck 
everywhere seal-brown ; on the sides of the neck the seal-brown 
colour is i-estricted to the extreme base of the hairs or, above the 
neck-glands, quite obliterated. — Occiput, crown, interocular space, 
and temporal region bright ochraceous-buff Avith a tinge of buff- 
yellow ; base of hairs (concealed on uppcrside of head, slightly 
shoAving through in temporal region) seal-brown. Muzzle, super- 
ciliaries, chin, and throat blackish. 

The coloration, it will be noticed from the above, is in every 
essential respect like that of Pt. comorensis and allied species, save 
for the very strong admixture of broccoli-brown and Avood-brown iu 
the colour of the back and rump. 

Size. — Much smaller than Pt. comorensis. 
3Ieasurements. On pp. 221, 222. 

Specimen e.vamined. One skin with skull, from the collection of 
the U.S. National Museum (Keg. no. 620G1). 
Ranr/e. Aldabra Island, N. of Comoro group. 
Colypes in the U.S. National Museum (nos. 20984, '85). 
Literature. — A few examples collected by Dr. W. L. Abbott, and 
one skull (Ji^i. s. c.) found by Dr. A. Voeltzkow in the house in 
Aldabra formerly occupied by Abbott, seem to be the only specimens 
recorded in literature. 


32. Pteropus niger, Kerr. 
Pteropus vulgaris, Dobson, Cat. Chir. B. M. p. 23. 

Vespertilio injrens, Clusiux, E.rot. libri decern, p. 94, c. %. (animal) 

(160-5 : Mauritius). 
Pteropu3 rufus aut niger, auviculis breribus acutiusculis : La Roiis- 

sette, L'risson, Regn. Anim. p. 216 (1756: Reunion) ; id.,op.cit. 

2 ed. p. 153 (1762: Reunion ). 
Le Chien-volant, Duubentun, Mem. Acad. R. Sci. Paris, p. 384 

(1759 : Reunion). 
La Roussette, Bnffon, Hist. Nat. x. pp. 55, 66, 81, pis. xiv., xv., 

xvi. (animal; tongue; skeleton) (1763: Reunion); id., op. cit. 

Suppl. iii. pp. 253-262 (1776: liabits) ; Fvuchet d'ObsunvtUe, 

JEs.tais pliihsojyhiques, p. 77 (1783: Reunion); i?. Geoffroy,Ann. 

Mus. a' Hist. Nat. vii. p. 227 (1806: Mauritius; habits in 

captivity) ; Milbert, Vuy. pittor. lle-de-F ranee, ii. p. 244 (1812 : 

Great Bat of Madagascar {nee Edtcards), Grant, Hist. Mauritius, 

p. 65 (1801 : Mauritius\ 
Vespertilio vanipvrus, Li7m. SipH. Nat. 10 ed. i. p. 31 (pt.) (1758) ; 

id., op. cit. 12 ed. i. p. 46 (pt.) (1766); P. L. 8. Miilter, Volht. 

Naturst/st. i. p. 152 (pt.) (1773). Var. A, Sckreber, Siiitg. i. 

pp. 153, 155, pi. xliv. (animal, copy from Buftbn) (1774). 

Boddaert, JElench. Anim. i. p. 68 (pt.; (1785). Var. a, Gineli?i, 

Linn. Si/st. Nat. i. p. 45 (1787) ; G. Cuvier, Tabl. Elem. p. 104 

(1798). Var. 2, Turton, Linn. 8t/.<^t. Nat. i. p. 24 (1802). 
Ptercipus vampyrus, var. n, Er.vleben, Si/st. Rer/n. Atiim. p. 133 

(1777); Desmarest, N. Diet. d'Hist. Nat. xix. p. 643 (1803: 

Spectrum vampirus, Lac^pede, Tabl. Mamm. p. 15 (1799) ; id., 

Mem. List. Paris, ui. p. 499 (1801); Daudin, in Luffon, Hist. 

Nat., Didut ed., Quudr. xiv. p. 188 (1802*). 
Pteropus (Spectrum) vamp\TUs, Matsehie, JfcTei'/ac'/uV. p. 29 (1899 : 

Mauritius); Troitessart, Cut. Mamm., Suppl. p. 54 (1904: 

Tamatave ; Reunion ; Mauritius). 
Vespertilio vampirus niger, Kerr, Anim. Kingd, i. pt. i. pp. xvii, fiO, 

no. 106 (1792). 
Pteropus niger, G. Fischer, Zorirjnosia, iii. p. 553 (1814). 
Vespertilio caniniis, Blumenbaeh, Handb. Naturt/. 5 ed. p. 73 (pt.) 

(1797) ; Goldfuss, Vergl. Naturbvschr. Sliur/. p. 98, pi. xi. (1809). 
Pteropus fuscus, E. Geoffroi/, Cat. Mamm. Mas. Nation. d'Hist. Nat. 

p. 46(1803: Reunion; Madagascar). 
Vespertilio niauritianus, Hermann (e Commer.ion 3LS.), Ohs. Zool.. 

p. 19 (1804 : ^lauritius) ; Oken. Lehrb. Naturg. iii. Abth. ii. 

p. 934(1816). 
Pteropu.s mauritianus, Schinz, Thierr. i. p. 155 (1821 : Mauritius). 
Spectrum vulgare, var. mauritiauuni, Gray, Cat. Monk. i^-c. p. 101 

(1870: Mauritius). 
Pteropus rufus {nee E. Geoff.), Tiedemann, Zooloyie, i. p. 535 (1808). 
Pteropus vulgaris, E. Geoffmy, Ann. Mii-s. d'Jlist. Nat. xv. p. 9l 

(1810: Reunion ; Mauritius) ; O/wu, Lehrb. Naturi/. iii. Abth.ii. 

p. 936 (1810); G. Cuvier, Rbym Anim. i. p. 124 (l817 : Mauri- 
tius; Reunion); Destnarest, Mamm. i. p. 109, no. 139(1820: 

* Titlepage dated 1799. On the true year of publication see C. W. Elck- 
mond, The Auk, xvi. no. 4, p. 329 (Oct. 1899). 

21 ti PiERurUS NIGER. 

Mascarenes) ; Schiftz, Thierr. i. p. 154 (1821: Mauritius; 
Reunion) ; Temminck, Man. Mamm. i. p. 182 (1825 : Mauritius ; 
Reunion ; ? Madagascar ; ? Afiica) ; Lesson, Man. Mnmm. 
p. 109, no. 279 (1827 : Mauritius ; Reunion) ; Gray, GnfUh's 
Aiiim. Kingd. v. p. 65, no. 155 (1827 : Mauritius ; Reunion) ; 
Desmarest, Diet. Sci. Nat. xlvi. p. 361 (1827 : Mauritius ; 
Reunion) ; Is. Geoffroy, Diet. Class. d'Hist. Nat. xiv. p. 699 
(1826 : Mauritius ; Reunion) ; J. B. Fischer, Syn. Mamm. p. 83, 
no. 7 (1829: Reunion; Mauritius; ? Madagascar) ; Wagler, 
Syst. Amph. p. 9 (1830) ; Temminck, Mun. Mamni. ii. p. 74, 
pi. xxxviii. (wliole fifj. of juv., sluill of juv. and ad.) (1837) ; 
Gray, Mag. Zool. ^ Bot. ii. p. 503 (1838: Mauritius; Reunion) ; 
Oken, Ally. Natury. vii. Abtli. ii. p. 985 (1838; Mauritius; 
Reunion); Wagner, Sc/ireber's Scmg., Sitppl. i. p. 350 (1839: 
Mauritius ; Reunion ; ? Madagascar ; ? Africa) ; Lesson, N. Tabl. 
R. An., Mamm. p. 13, no. 177 (1842: Reunion; Mauritius); 
Schinz, Syst. Verz. Siiug. i. 'p. 124 (1844: Mauritius; Reunion) ; 
Wagner, Schreber's Sdvg., Siippl. v. p. 602 (1853-55 : Mauritius; 
. Reunion) ; Gervais, Hist. Nat. Mamm. i. p. 188, iig. p. 189 

(animal) (1854 : Mauritius ; Reunion ; Madagascar) ; Giebel, 
Snug. p. 997 (1855 : Mauritius ; Reunion) ; Schlegel, Dierkiinde, 
i. p. 63 (1857 : Reunion ; Mauritius) ; Peters, MB. Akad. 
Berlin, 1867, p. 323 (Mascarenes) ; Fitzinyer, SB. Akad. Wien, 
Ix. Abth. i. p. 449 (1870: Reunion; Mauritius); Bartlett, 
P. Z. S. 1875, p. 03 (Tamatave) ; Dobson, Cat. Chir. B. M. p. 23 
(1878 : Mauritius) ; Troue.ssart, Bev. S,- Mag. Zool. (3) vi. p. 204 
(1879: Tamatave; Reunion; Mauritius); Mobius, Meeresfauna 
Mauritius,T^.uQ (ISQO: Mauritius; habits); Jentink, Cat. Ost. 
ilfa»w?n. p. 252 (1887: Mauritius; Madagascar); id., Cat. Syst. 
Mamm. p. 138 (1888 : Mauritius ; Madagascar) ; H. Allen, Pi-oc. 
Ac. Nat. Sci. Philad. 1 889, p. 336 (wing-membranes) ; Trotiessart, 
Cat. Mamm. i. p. 77 (1897 : Tamatave ; Reunion ; Mauritius) ; 
Matschie, Megachir. pi. ix. figs. 2, 2 a, 2 b (skull) (1809). 

Spectrum vulgare, Gray, Cat. Monk. ^c. p. 100 (1870: Mauritius; 
Reunion : ? Madagascar). 

Pteropus edwardsi (pt., ncc E. Geoff.), Pollen, in Pollen ^- v. Dam, 
Eech. Faune Madagascar, ii. p. 26 (1868: Reunion). 

Pteropus pteropus, " Brisson,^' Merriam, Science, (n. s.) i. no. 14, 
p. 376 (5 Apr. 1895). 

Diac/nosis, — Skull and dentition (fig. 12) quite as in Pt. rufus and 
comorcnsis. Ears small, almost concealed in the fur. Tibia thickly 
haired above. Sides of back and rump buffy, contrasting with 
dark spinal tract. Forearm 159-171 mm. Hab. Mascarenes. 

Ears. — Small, almost hidden in the long fur. Tip sharply 
pointed, outer margin distinctly concave below tip. Hinder face 
of conch clothed with short, thinly-set hairs on basal half, tip 

Interfemoral.— Centred portion concealed by the far, scarcely 
more than 2 mm. in depth. 

Far. — Longer than in Pt. rufus ; directed backward but not 
closely adpressed on back, semierect on nape of neck. Approximate 
length of hairs at middle cf back 26 mm., middle of mantle 27 mm., 
middle of belly 26 mm. Width of furred area at middle of back 
about 65 mm. 



Above, humerus and proximal third of forearm clothed with 
short, densel5'-set fur; femur and tibia to a short distance from 
ankle long-haired ; small bunches of rather long hairs on upperside 
of foot at base of first phalanges and distal extremity of second 
phalanges. Eelow, humerus and proximal part of forearm and 
tibia densely furred ; closely-set woolly fur on membrane along 
outer side of forearm, and on plagiopatagium to a line between 
elbow and knee. 

Fig. 12.— Fieropus niger, (S ■ Mauritius. No. }. 

Colour. — Chief characters : face, sides of back and rump, anal 
region, and fur on tibia above and below huffy, strongly contrasting 
with dark mantle, spinal tract, and underparts. 


Mauritius specimens : — Median tract of buck and rump, from 
ehoulders to interfemoral, approximately vandj'ck-brown, darkest 
(tinged with seal-brown) anteriorl}-, becoming gradually lighter 
(more pure vandyck-broAvn) posteriorly; the width of this dark 
longitudinal area varies individually between 28 and 38 mm. 
Sides of back, rump, and fur on upperside of tibia buff, strongly 
contrasting with dark spinal region. — Breast, belly as far back as 
anal region, and flanks dark seal-brown with a peculiar oily gloss, 
and slightly washed with mars-brown on middle of breast and 
belly ; anal region and fur on underside of tibia buify or ochraceous- 
buft'y. — Mantle glossy blackish seal-brown, darkest in the middle, 
passing posteriorly gradually into the dark colour of the spinal 
stripe, anteriorly tinged with chestnut, aud lightening on occiput 
to dark cinnamon-rufous ; sides of neck similar to mantle, shading 
into golden tawny posteriorly near insertion of antebrachial 
membrane ; foreneck blackish seal-brown, similar to (or rather 
darker than) breast. — Interocular space and sides of face huffy or 
ochraceous-bulfy ; crown, temporal region, chin, and throat bufty 
more or less strongly tinged with tawny or pale hazel. 
Measurements. On pp. 221, 222. 

Sjjecimens examined. Ten, in the collections of the Paris, Leyden, 
Berlin, and British Museums. 

Mange. The Mascarenes : lleunion, Mauritius. 
Some early writers have recorded this species from Madagascar, 
as a rule, however, with a sign of query. The only recent records 
from that island known to me are Bartlett's in P. Z. S. 1875, p. G3, 
of a specimen stated to have been obtained at Tamatave (bv 
Mr. Waters), and Jentink's (Cat. Ost. p. 252 ; Cat. Syst. p. 138) of 
a specimen from Madagascar (Mulie, 1876). The former I have 
not seen ; the Leyden specimen does not ditler from Mauritius 
examples. The correctness of the locality is in both cases open to 
doubt ; the specimens may have been brought alive from the 
Mascarenes to Madagascar. 
Tj/2)e not in existence. 

Hahits. — This species, de la Nux wrote from Eeunion in a letter 
to Buffon (Suppl. iii. pp. 253-267), is only found in the forests ; it 
rests in the crowns of the trees, suspended from the branches, 
wrapped in its wings, not (as Pt. suhniger) in hollow trees or caves. 
It is less strictly nocturnal in its habits than Pt. suhniger ; by day- 
time it is sometimes seen flying at a considerable height, and it is 
probable that it not infrequently passes from Reunion to Mauritius 
and vice versa. Its favourite food in Eeunion is bananas, peaches, 
guava, mistletoe-berries, &c., but the food varies of course greatly 
according to the season ; also it is fond of sucking the juice of the 
blossoms of various species of Umbellifers ; at the time of the year 
(mid-summer, i. e. January-February) these plants are in bloom, 
the lioussettes come in great numbers down to the lower j)arts of 
the island, and the ground may here and there be found literally 
covered with blossoms torn off by these bats. Pairing takes 
place about the middle of autumn (May) ; the period of gestation 


ia four and a half or five months, so that the single young 
is generall)' born about one month after spring equinox (October), 
After eight months (winter solstice, about June) the young is full- 
grown. In 1722, on his arrival at Keuniou, de la Nux found the 
Koussettes extremely common ; abundance of food -would attract 
flocks of a hundred, a hundred and fifty, or even two hundred to 
one spot ; during his more than fifty years of residence in the island 
the number of individuals rapidly decreased, chiefly owing to de- 
forestation of the island and the shooting of these Fruit-bats for 

Earliest records of the species. — Few, if any, species of Fruit-bat 
can be traced so far back in literature as Pt. niger. The example 
named by Lecluse (1G05) Vespertilio inrjens was seen by him in 
Amsterdam in 1G03, and stated to have been brought from llha do 
Cerne (Mauritius) ; his woodcut alone would almost be sufficient 
for a safe identification of the species ; the description and locality 
exclude all doubt. 

The habitat of the species accounts for the fact that it was 
particularly well known to the French zoologists in the latter half 
of the eighteenth century. It is Daubenton's " Chien-volant" from 
Eeunion (1759), of which he gives an excellent detailed description. 
Whether his " Roussette " from the same island (Z. s. c. p. 385), 
stated to differ from the Chien-volant in colour only, but of which 
he had only seen one specimen in bad condition, is Pt. nir/er or 
Pt. suhniijer is open to question. 

Brisson (1756 and 1762) discriminated three species of Pterojnis: 
Pt. ritfus ant niger., " La iloussette," from Reunion, which is un- 
questionably /*<. ?u(/^?*; but with this species Brisson erroucously 
united Seba's Canis volans Ternatamis Orievfalis ; further, Pt. collo 
riihro, from Reunion, which is Pi. suhniger ; and Pt. auricidis 
'patulis, from Nova Hispania, which is Vampynim spectrum. 

Buffon (1763) distinguished sharply between La Roussette and 
La Rougette, both from Reunion ; the former is Pt. niger, the latter 
Pt. suhniger. In Buffon's time the Roussette was represented in 
the Royal Cabinet by one specimen, mounted with expanded wings, 
sent from Reunion by de la Nux, one skeleton, and some anatomical 
preparations ; Buffon's plate shows the light-coloured sides of the 
back so characteristic of Pt. niger. No later writer has given a 
better, more detailed and vivid account of the habits of this species 
than de la Nux (quoted by Buffon, 1776). Buftbn's specimens are 
probably no longer in existence ; none of the specimens I have 
seen in the Paris Museum can be identified with those described by 

Vespertilio (and Pteropus) vampgrns, of Linne and the early 
post-Linnean writers. — Linne's V. vampyrxts (1758 and 1766) is a 
mixture of three species, viz. Pt. vampyrns (i. e. " Pt. edulis, Geoff.," 
as understood by Dobson), based on Scba ; Pt. niger, based on 
Lecluse ; and Pf. rvfvs, based on Edwards (Great Bat from 
Madagascar). Techuically the name vampyrus must stand for the 
first of these species. 


Most of the earlier post-Liunean writers (Schreber, Erxlebcn, 
Boddaert, Gmelin, Kerr, &c.) divide " vampyrus " into three varie- 
ties : a larger, which is Pi. niyer, or a mixture of Pt. nif/er aud 
Pt. vampyrus, sometimes also Pt. rufus ; a smaller {Pt. subniyer) ; 
and a straw-yellow {Eidolon helvum). Schreber, Erxleben, and 
Gmelin designated these varieties by letters only (A, B, C, or a, /3, 
-y) ; Kerr was apparently the first to distinguish them by technical 
names (niger, suhntger, helvus). 

VespertiUo vampirus niger, Kerr : 1792. — Based by Kerr on 
Schreber's figure (vol. i. pi. xliv.) of V. vampip-us var. A, which is 
a copy of Buffon's plate (vol. x. pi. xiv.) of La Roussette. The type 
locality of this latter, viz. lleunion, must, therefore, be fixed as the 
type locality of V. v, niger. Most of the synonyms given by Kerr 
under the heading V. v. niger (Clusius, Brisson, Daubenton) also have 
reference to the present species ; the rest partly (Seba, Pennant) 
to Pt. vampyrus, partly (Boutins) to a Flying Lemur. 

VespertiUo caninus, Blumenbach ; 1797. — Name proposed in 
lieu of the Linnean V. vampyrus, because this bat " blosz von 
Baumfriichten lebt und also schlechterdings nicht Vampyr genannt 
werdeu kann." Diagnosis a verbal copy from Linne ; references 
three, viz. Linne's vampyrus [i. e. Pt. vampyrus, niger, and r«/«.s], 
Bufl'on's lloussette [P<. niger'], and Schreber's pi. xliv. [Pt. niger], 
— Goldfuss's plate of " V. caninus Blumenb." (1809, 1. s. c.) is a 
copy of Schreher's plate xliv., of V. vampyrus var. A, i. e. Pt. niger. 

Pteropus fuscus, E. Geoffrey ; 1803. — Type locality, lleunion. 
The following statements in the diagnosis are sufficient for an 
identification of the species : " des polls roux [in the description 
' roux-jaunatres '] sur la face, autour de I'anus, et sur les parties 
laterales du dos." Based by Geoffro}- on three specimens in the 
collection of the " Museum national d'histoire naturelle," viz., two 
mounted specimens (wings expanded), sent from lleunion by 
de la Nux (nos. 86 and 87), and a third " envoye de Madagascar, 
par le citoyen Mace, naturaliste " (no. 88) ; in addition, the Museum 
possessed some embryos in alcohol (no. 89). As mentioned above, 
Buffon's specimen of the Roussette (mounted, wings expanded ; 
ancient Royal Cabinet) was obtained in Reunion by de la Nux ; 
since also two of Geoffrey's cotypes of Pt. fuscus (both mounted, 
wings expanded) had been sent from Reunion by de la Nux, it is 
possible that the former specimen was identical with one (no. 86 
or 87) of the latter. Geoffrey's specimen no. 88, stated to have 
been sent from Madagascar, was probably a dealer's specimen ; the 
fact that Geoffroy in 1810, on redescribing Pt. fuscus under the 
name Pt. vulgaris, gave as habitat only '' I'ile de France et celle 
de Bourbon," without any reference to the Madagascar specimen, 
may indicate that he was himself in doubt as to the correctness of 
this locality. None of the s})ecimens I have seen in the Paris 
Museum can be identified with Geoffrey's cotypes of Pt. fiiscus. 

VespertiUo mauritianus, Hermann ; 1804. — Type localit}', Mau- 
ritius. Based on MS. notes by Commerson. The words " capite 
dorsique lateribus rufis," together with the locality, exclude all 
doubt as to the identification. 



Ftcrojjus rvfus, Tiederuanii; 1808.- — Based on Brisson's Ft. rufus 
aut niger (the name Ft. rufus is evidently an abbreviation of 
" Ft. rujus aut niger ") ; other synonyms quoted by Tiedemann : 
Vesp. vamjjyrus, L., which is partly Ft. niger ; Vesj). caninus, 
Blumenb., BufFon's lioussette, and Schreber's pi. xliv., all of which 
are Ft. niger. Name preoccupied by Ft. rufus, E. Geoff., 1803, 
which is Ft. ediuardsi, auctorum. 

Fleropus vulgaris, E. Geoffrey ; 1810. — A redescription and 
renaming of Geoffrey's Ft. fuscus, 1803 (but without any reference 
to this latter, because Geoffroy wished to suppress his Catalogue 
from 1803). The type locality ot fuscus, viz. Beunion, must there- 
fore be fixed as the type locality of vulgaris, too ; it is even 
highly probable that the latter was based on the same individuals 

External measurements of Pteropus seychellensis, 
aldabrensis, and niger. 



6 yg. ad. 

(Incl. cotypes.) 


Pollex, total length, c. u 

„ metacarpal 

„ Ist phalanx 

I 2nd digit, metacarpal 

„ Ist phalanx 

,. 2nd-3rd plialanx, c. u. 

! 3rd digit, metacarpal 

i ,, Istphalanx 

\ „ 2nd phalanx 

4th digit, metacavpal 

., 1st phalanx 

„ 2nd phalanx 

5lh digit, metacarpal 

„ ] st phalanx 

„ 2nd phalanx 

Ear, length from orifice 

,, greatest width, llattened .... 
Interfemoral in centre, depth.... 

Lower leg 

Foot, c. u 





























































Pt. niger. 
2 ad. 

MiN. Max. 

mra. mm. 

159 171 

69 72-5 























60-6 64 




* The four minimum measurements of Pt. seychellensis marked with an 
asterisk are taken from specimens which I suspect to be not quite full-grown ; 
the other examples measured, though also slightly immature, are probably 

t E.stimate from dried skin (in the two cotypes, which also are skins, the 
ears measure, according to True. 24 and 25 mm.). 


Measurements of si' alls and teeth of Pteropus seychellensis, 
aldabreusis, mid niger. 



$ imni. 





Pt. niger. 
Skull and 
teeth: 1 ad. 

Skull, total length to gnatliion 


























































































,, palation to incisive foramina ... 
„ front of orbit to tip of nasals ... 
,, width of brain-case at zygomata 
,, zygomatic width 

„ width across m', externally 

„ lachrvinal width 

„ width across canines, externally. 
,, postorbital constriction 

„ interorbital constriction 

„ width of mesopterygoid fossa ... 
„ between p''-p', internally 

,, between cingula of canines 

„ orbital diameter 

Mandible, length 

„ coronoid height 

Upper teeth, c-m^ 

Lower teeth, c-m, 

Upper incisors, combined width 

p^, length 

,, widtli 

pS length 

„ width 

m^, length 

,, width 

m"', length 

,, width 

p.. length 

,, width 

„ width 

P4, length 


as the former. — After a brief description of Pt. vulgaris, Geoffroy 
refers to " un autre individu, deniierement apporte de I'ile de 
France," and which he regards as " une variete dans cette espece" 
ou account of its slightly different colour. Q'he Paris iluseum 
possesses a specimen (mounted, wings expanded, skull in situ ; 
recent Register no. A. 1) marked in handwriting on the underside 
of the block " Pterojms vulgaris (Geotf.) ; type " (no locality 
written), and on the printed label " Ancien Cabinet ; Maurice." 
This specimen, which certainly bears the appearance of having 
been preserved for many years in the Museum (much bleached, back 
naked), is, most probably, not one of the true cotj^pes of Pt. vulgaris 
(and Pt. fuscus), which were obtained in Reunion, but Geoffrey's 
" variety " of Pt. vulgaris referred to by him in the words quoted 
above ; if so, it is the individual mentioned by Geoffroy in 1806 
(Ann. Mus. d'Hist. Nat. vii. pp. 227-230) as having been kept in 
captivity by Surgeon-Major Eoch, Mauritius, and brought by him 
to France in 1803. 

" Pteropus pteropus, Brisson," Merriam ; 1895. — Combination 
introduced into literature by Merriam on the mistaken supposition 
that Brisson's nomenclature was binomial. 

RemurTcs. — Pt. nigcr is readily discriminated from any other 
species of the genus by the colour of the upperside : pale sides of 
back contrasting with dark mantle and spinal tract. 

a. Yg. ad. st. Pui-.-hased (Gardiner). 

b. 6 yg- ad. sk. ; skull. Mauritius. H. Whitely, Esq. [P.]. 

E. The Pteropus melanotus group. 

Species. — Pt. melanotus, tytleri, niadicus, modiglianii, natalis. 

liange. — Nicobar Islands, north to the Audamans, south through 
Nias and Engano, to Christmas Island (S. of .Java). — This is the 
absolutely dominant (and probably truly indigenous) type of 
the genus in the Andaman-Nicobar chain, tlie only other groups 
represented being the widely spread liypomelanus and the Indo- 
Malayan vampyrus groups. 

Qenerul characters. — Skull typical Pteropine. Dentition without 
special modifications ; cingulum of canines generally rather narrow ; 
posterior basal ledges of p\ Pg, and Pj short, but distinctly marked off 
from teeth. Ears rather large, broad, scarcely attenuated above, 
tip rounded off ; interfemoral generally distinctly developed in 
centre (obsolete in Pt. natalis) ; tibia practically naked above. 
Prevailing colour of fur blackish above and beneath, either entirely 
uniform, or with bright colour restricted to mantle or to mantle 
and centre of breast and belly ; blackish colour in one species 
(Pt. niadicus) varied with greyish. Sexual differentiation incon- 
spicuous (as in hy pomelamis group). Size moderate or rather large 
(forearm 125-165 mm.). The extreme south-eastern species 
{Pt. natalis) is slightly aberrant in dentition. 

Specific dlfftrentiation. — The Nicobar species (Ft. melanotus) is 
apparently the least modified form of the group: fur short, bright 

224 PTERorrs melanotcs. 

mantle aud centre of breast and belly contrasting with general 
dark colour of pelage. The Andaman species (Pt. tytleri) is either 
entirely blackish or with some indication of a brighter tinge in the 
mantle ; general size smaller. The Nias species {Pt. niadicus) is 
in all essential characters similar to Pt. melanotus, but with a con- 
spicuous admixture of greyish in the dark-coloured parts of the fur. 
In the Engano species (Pt. modigliann) the fur is longer and uniform 
blackish, the size smaller than Pt. melanotus. In the extreme 
south-eastern species, Pt. natalis (Christmas Island), the dentition 
is somewhat modified (see p. 233) ; as in Pt. modiglianii the colour 
is uniform blackish, with or without a faint trace of a paler tippet, 
the fur still longer, the size still smaller. 

AJJinities of group. — The Pt. melanotus is undoubtedly closely 
related to the Malagasy Pt. rufus group, with the typical members 
of which it accords in all essential characters of the skull and 
dentition, distribution of fur, and development of interfemoral ; it 
differs chiefly in the shape of the ears and a pronounced tendency 
to suppression of all light colours. The five species of the melanotus 
group are so nearly interrelated that they may be presumed to be 
insular modifications of one type of bat which inhabited a land area 
of which the Andamans, Nicobars, Simalu, Nias, Mentawei, Engano, 
and Christmas Island are probably the remnants. 

33. Pteropiis melanotus, Blyth. 
Pteropus nicobaricHS (pt.), Dobson, Cat. Chir. B. M. p. 54. 

Pteropus edulis {nee E. Geoff.), Blyth, J. A. S. B. xv. p. 367 
(1846: Nicobars) ; Peters, MB. Akad. Berlin, 1867, p. 324 (pt.) 

I'teropus melanotus, Blyth, Cat. Mamm. Mus. As. Soc. p. 20 (1863: 
Nicobars) ; id., Mouat's Andaman Islanders, App. p. 354 (1863 : 
Nicobars) ; Mason, Bee. hid. Mus. ii. pt. ii. p. 159 (1908 : 

Pteropus nicobaricus, Fitzinyer, SB. Akad. Wien, xlii. p. 389 (1861 : 
Nicobars) (nom. nud.) ; Blyth, Mouafs Andaman Islanders, App. 
p. 354 (1863: Nicobars) (liom. nud.); Zelebor, Reise ' Novara,'' 
Zool. i. Sdug. p. 11 (1869: Car Nicobar) (descr. princeps) ; 
Fitzinger, SB. Akad. Wien, Ix. Abth. i. p. 410 (1870 : Car 
Nicobar) ; Dohson, J. A. S. B. xlii. pt. 2, p. 198 (pt), pi. xiv. 
fig. 2 (ear) (1873 : Nicobars) ; id., P. Z. S. 1873, p. 250 (pt.j 
(Nicobars ; sexual colour difference) ; id., Cat. Chir. Ind. Mus. 
' pp. 2-3 (pt.) (1874: Nicobars); id., Journ. Anthrop. Inst. iv. 

pt. 2, p. 458, footnote (pt.) (1875 : Nicobars) ; id., Mon. As. 
Chir. p. 17 (pt.) c.fig. (head), pp. 22, 188 (1876: Nicobars) ; id., 
Cat. Chir. B. M. p. 54 (pt.) (1878 : Nicobars) ; Trouessart, Bev. 
tV Mag. Zool. (3) vi. p. 202 (pt.) (1879: Nicobars) ; J. Anderson, 
Cat. Mamm. Ind. Mus. pt. i. p. 102 (pt.) (1881 : Nicobars) ; 
Jentink, Cut. Syst. Mamm. p. 147 (pt.) (1888: Katchall) ; Blan- 
ford, Fauna Brit. Ind., Ma7nm. pt. ii. p. 260 (pt.) (1891 : 
'Nicobars): Lydekker, R. Nat. Hist. i. p. 256 (pt.) (1893-94); 
Trouessart, Cat. Mamtn. i. p. 81 (pt.) (1897 : Nicobars) ; 
Matschie, Megachir. p. 16 (pt.) (1899: Nicobars); Miller, 
Proc. U.S. Nat. Mus. xxiv. p. 782 (1902 : Great Nicobar : Tillan- 

rTi:uoi'Us MELixoxcs. 225 

chon;;) ; Kio)i!<, Anduinans ^- yicobars, pp. 70, 13;3, 82-5 (1903: 
(Jar Nicobar ; Tillanclioug; Great Nicobar ; habits); Trouessart, 
Cat. Mamm., Suppl. p. 50 (pt.) (1904 : Xicobars) ; Miller, Fam. 
8f Gen. Bats, p. 58 (pt.) (1907). 

Diagnosis. — Skull typical Pteropine. Posterior basal ledges of 
p', P3, and p^ distinctly marked oaf from teeth. Ears large, broad, 
exposed, tip rounded off. Tibia naked above. Fur of back short, 
adpressed. Mantle and centre of breast and belly bright-coloured, 
contrasting with blackish back, head, throat, and flanks, Si/.o 
above medium : forearm 153-16-3 mm. Hah. Nicobar Is. 

Skull. — As in Pt. riifus. 

Teeth. — Essential characters as in Pt. ritfas. Cingulum of upper 
canines rather narrower, p^, m\ and p^ somewhat heavier, m, 
smaller. Posterior basal ledge of ]f short, but distinctly marked 
off from tooth postero-externally ; in p^ the ledge is usually more 
developed postero-internally than postero-externally, making pos- 
terior margin of crown more or less oblique on longitudinal axis of 
tooth. Posterior ledge moderate in p^, short in Pj, in both teeth 
distinctly separated postero-externally from base of outer main cusj). 

Palate-ridges. — 5-|-5-l-3. First ridge terminating laterally at 
front of canine; second at back of canine; third at front of p^ ; 
fourth at back of p^ ; fifth at back of p' ; sixth at front of m^ ; 
seventh at front of m" ; eighth to tenth behind m" ; eleventh to 
thirteenth situated near palation border. 

Ears. — -Kather large, broad, exposed. Inner margin stronglv 
and evenly convex from base to tip ; outer margin strongly convex 
in lower two-thirds, concave in upper third ; tip rather broadly 
rounded off. Naked, except posteriorly at base and along basal 
half of outer and inner margins. 

Wings. — Membranes arising close together near sides of spine 
(interspace about 10-17 mm.). 

Inter femoral. — Much reduced in centre, depth about 2-4 mm. 

Fur. — Short ; straight and adpressed on back ; wavy, somewhat 
crinkled and spreading on rump and mantle. Longest hairs of 
back and mantle about lo-lG, of belly 13-14 mm. Width 
of furred area of back about 50-55 mm. 

Above, a narrow line of adpressed hairs along upperside of 
humerus. Forearm naked, except for some very thinly spread and 
closely adpressed hairs. Knee and tibia practically naked. Inter- 
femoral naked, except in central portion. — Below, forearm, tibia, 
and lateral interfemoral naked. Woolly hairs on antebrachial 
intmbrane, along outer side of forearm, and on lateral membrane 
next to body. 

Colour. — Whole scries examined, adult males and females, skina : 
Back and rump generally blackish seal-brown, occasionally witli a 
slight tinge of dark vandyck-brown. A few greyish-white hairs as 
a rule detectable on close examination.- — -Middle of breast and belly 
bright-coloured, contrasting with blackish sides, flanks, and anal 
region. Bright-coloured area varying individually from golden 
ochraceoua-bulf (^lightest extreme) to rich tawny, along the median 


226 PTERorcs melanotus. 

line of breast and belly sometimes tinged with pale drab; laterall)', 
before merging into the blackish of the sides, this bright colour 
generally shades into eiunamon-rufous or russet. Concealed base 
of bright-coloured hairs blackish seal-brown.^ — Mantle much like 
bright centre of breast and belly, varying individually from pale 
ochraceous-buff, through tawny, to almost cinnamon-rufous ; even 
the palest extreme (pale ochraceous-buff) is generally slightly 
clouded with orangc-buiT, and shades posteriorly and anteriorly 
(before merging into blackish back and blackish head) and laterally 
(sides of neck) into tawny or cinnamon-rufous ; the dai'kest extreme 
is almost cinnamon-rufous in the centre, shading in front, behind, 
and on sides of neck into hazel or chestnut. No (or only ex- 
tremely short) blackish bases to bright-coloured hairs. Foreneck 
russet, often slightly mixed with, occasionally almost obscured by, 
blackish hairs. — Crown, as far as liack of ears, generally blackish, 
in strong contrast to bright mantle, but occasionally more or less 
strongl}' tinged with russet. Sides of head, chin, and throat 
blackish or seal-brown. 

No appreciable sexual difference in colour (six males, three 
females, one skin of doubtful sex examined). 

Measurements. On pp. 230, 231. 

Specimens examined. Ten, from the collections of the Yienna 
(one, cotype of Pt. incoharkits), U.S. National (five *), and Eritish 

lianf/e. Nicobar Islands, the whole group. Specimens examined 
from Car Nicobar, Tillanchong, Trinkut, and Great Nicobar. 

CoUfpes in the Calcutta Museum. 

Habits. — In Great Nicobar, early in March, 1901, Dr. W. L. 
Abbott and Mr. C. B. Kloss found a colony of these bats, number- 
ing several thousands, in mangroves on either side of a small river. 
The surrounding atmosphere was impregnated with the musky 
odour of their bodies. When disturbed they gave vent to a con- 
tinuous " skirling " noise, somewhat like the song of cicadas, but 
less shrill in tone. By nature they were very fearless, and the 
majority merely stared inquisitively; a few spread their Avings and 
flapped heavily away for a short distance, while others crawled 
actively along the branches back downwards. All the females 
carried, clinging to the breast, a young one of about one-third full 
growth ; these the mothers hugged to themselves with a folded 
wing, but when unsupported, the young found no difficulty in 
maintaining its position, by means of its claws and its suction grip 
on the ])arent's teat. When the latter crawled about the body was 
supported in the membrane of the wing, which bagged slightly 
'with the weight. The action of these bats when climbing a vertical 
branch is similar to a man's in climbing up a pole. The wings are 
first raised and a tight grip taken with the claw of the thumb, 
then the feet are drawn up, and, after they obtain a hold, the 

* U.S. N, ]\r. nos. 111729 ami 111732 (Tillauciiong), 111738-10 (Great 

rn:it(ii'us Tvn.KRr. 227 

wings aro onco moro lifted. When taking to llight,, they swing to 
and fro once or twice, and then let go in a backward direction. 

Pteropxis melanotus, lilyth ; 1863. — Three examples of this 
species, collected in the " Nicobar Islands " by Capt. Lewis and 
presented by him to the Calcutta Museum, were briefly described 
(colour of fur) by Blyth, in 1846 (/. s. c), wlio referred them to 
" Pi. edulis : Ft. javanicus, Horsf., &c., &c.," though with some 
em})hasis on the peculiarities of the specimens. Seventeen years 
later, Blyth (1863, 7. s. c.) proposed for the same specimens (now 
reduced to the number of two) the name Pt. meliDwtus. Name 
evidently given in allusion to the blackish back (melnnonotiis), not 
to the black ears (meUuiohig). 

Fti'fopus nicohariciis, Zelebor ; 1869. — Type locality, Car Nico- 
bar. Two cotypes, cJ ad. skin, J jiin. al., Yienna Museum; I have 
exaiuined the latter and the skull of the former. 

«. 2 iium. al. ; Nicobars. Lidian Museum [E.l. Not reg. 


h. Ad. sk. ; [Nicobars.] A. O. Huine, Esq. [P.]. 


c, t/. c? ad., O ad. Trinkut, Nicobars, A. O. Hume, E?q. [P.]., 
ska.; skulls. 17 Feb. 1873 (jr. 108. 

'64. Pteropus tytleri, Mason. 
Pteropus nicoharinis (pt.), Dobson, Cat. Chir. 15. M. p. 54. 

Pteropus nicobaricus {nee Zelebor), Dobson, J. A. 8. B. xlii. pt. 2, 
p. 198 (pt.) (1873 : Andamans) ; id., P. Z. S. 1873, p. 2.jO (pt.) 
(Audamans ; sexual colour difference) ; id., Cat. Chir. Ltd. Mus. 
pp. 2-3 (pt.) (1874 : Andamaus; colour of mantle) ; Ilwnc, Stray 
Feathers, ii. p. 61 (1874 : Little Jolly Boy) : IJobson, Journ. 
Anthrop. Inst. iv. pt. 2, p. 458, footnote (pt.) (1875 : Andamans) ; 
id., Mon. As. Chir. pp. 17, 22, 188 (pt.) (1876: Andamans; 
colour of mantle); id., Cat. Chir. B. M. p. .54 (pt.) (1878: 
Ross I.) ; Trouessart, Rev. S,- Ma;/. Zoul. (3) vi. p. 202 (pt.) 
(1879 : Andamans) ; J. Awlerson, Cat. Mamm. hid. Mus. pt. i. 
p. 102 (pt.) (1881: Andamans); Jeiitink, Cat. Out. Alamm. 
p. 259 (ls87 : Audamans) ; id., Cat. l^yst. Mamm. p. 147 (pt.) 
(1888: Andamans); Blanford, Faun. Brit. Ind., Mamm. \it.\\. 
p. 2G0 (pt.) (1891: Andamans); Trouessart, Cat. Mamm. i. 
p. 81 (pt.) (1897: Andamans); Matschie, Meijachir. p. 16 (pt.) 
(1899: Andamans); Miller, Proc. U.S. Nat. Mus. xxiv. p. 782 
(pt.) (1902 : S. Andaman; Little Jolly Boy) ; Kloss, Andamans 
and Nicobars, p. 325 (pt.) (1903 : 8. Andaman ; Little Jolly 
Boy) ; Trouessart, Cut. Mamm., Sujipl. p. 50 (pt.) (1904 : 

l't(>ropus tvtleri, Dcbson, Cat. CJiir. Ind. Mm. p. 3 (1874 : S. Anda- 
man) (nom. uud.); id., Mon. As. Chir. p. 189 (1876: S. Anda- 
man) (nom. nud.) ; Mason. Per. Ind. .Mna. ii. pt. ii. p. 1('>2 (pt.) 
(July, 1908 : Rutland ; S. Andaman ; Little J. lly Boy). 

Dittf/nosis. — Similar to Pt. melanotus, but smaller, and with the 
whole of the undcrparts uniform blackish ; mantle often 



like back, occasionally tinged with a brighter colour. Forearm 
about 149-5 mm. Hah. Andaman Is. 

Slndl and teeth. — Skull as in Pt. melanotus, but conspicuously 
smaller ; mandible about 53 mm, against 56'5-(30 in the allied 
species. Dentition differing in no respect from that of the IV^icobar 
species, except perhaps in a slightly smaller average size of the 

Colour. — 5 ad., type, teeth much worn ; imm. (probably full- 
grown), sex indeterminable ( : — Uniform blackish or 
blackish seal-brown above and beneath, faintly tinged with dark 
vandyck-brown on rump ; a few greyish-white hairs among the 
dark ones on back and muz/le. Middle of breast and belly not 
differing from sides and flanks ; underparts not differing from back, 
except perhaps in the still more intensely blackish tinge of the 
colour ; mantle in certain lights tinged with seal-brown. 

A young male (rather more than half-grown ; is 
precisely similar to the adult female, excejjt for a rather more 
pronounced seal-brown colour of the mantle (with a faint sugges- 
ion of chocolate). 

According to Dobson (P. Z. S. 1873, Z. ,«. c.) females, from the 
Andamans and Nicobars, are " generally of an intensely black 
colour throughout ; in a few specimens only, of apparently very 
aged individuals, the fur on the back of the head and neck has a 
slightly reddish tinge ; while the males have the whole of the head 
and nape of the neck to the shoulders bright orange or pale yellow 
(very rarely, in old males, reddish brown) as in Pt, medius, con- 
trasting as remarkably with the sombre hues of the females as the 
brilliantly coloured skin of the male Mandrill contrasts with the 
same parts in the other sex." So far as Nicobar individuals 
(Pt. melanotus) are concerned, the alleged sexual difference in 
colour is imaginary ; all the specimens I have seen (ten), females 
as well as males, have the mantle, occiput, and middle of breast 
and belly bright-coloured, contrasting with the blackish colour of 
the rest of the pelage, and such variation in the bright tinge as 
does occur is individual, not sexual (compare also Miller, Proc. U.S. 
Jfat. Mus. xxiv. p. 784 ; 1902). As to the Andaman species 
(Pt. tytleri), the few specimens available, one of which is a male, 
another a female, and a third of doubtful sex, are uniform blackish 
with the mantle practically similar to the back ; but from Dobson's 
notes (1874 and 1876, I. .9. c.) on the colour of the mantle of some 
Andaman individuals in the Calcutta Museum it appears certain 
that in this as in several other melanistic species of Pteropus the 
mantle is occasionally tinged with a brighter colour. It is probable 
that Dobson, by comparing Andaman with Nicobar individuals, 
mistook what is really a specific difference between Pt. ti/tleri 
(practically uniform blackish) and Pt. melanotus (bright mantle 
and centre of breast and belly) for a seocual colour difference 
between individuals of one species (" Pt. nicobaricus ''). 

Measurements. On pp. 230, 231. 
■ Specimens ea-amined. The British Museum material. 


Ramie. Andamau Islands: Kuthuid Island, iSouth Audaman, 
Little Jolly ]3oy. 

l\ipe presumably in private possession. 

Ilistovy of technical name. — Originally a manuscript name written 
by Blyth on the labels of three skins, obtained in South Andaman 
Island by Lieut.-Col. Tytlcr and ])reseuted by him, in 1864, to the 
Calcutta Museum (specimens 22, 23, and 24 in Dobson's Mon. 
Asiat. Cbir. p. 18<S ; specimens d, e, and / in J. Anderson's Cat. 
Mamm. Ind. Mus. i. p. 102, specimen/ wrongly stated to be from 
S. Nicobars instead of S. Andamans). The name remained un- 
published until printed, without description, by Dobson in 1874 
and 1876 (7. s. c). The type of Mason's Ft. t)^tleri (1908, I. s. c.) 
is stated to be an adult male, skin and skull, Rutland I., collected 
by B. E. Osmaston, March 5, 1907. Mason failed to point out 
the real differential characters of the species as compared with 
Pt. melanotus. His statement that the two sexes differ in colour 
to such extent that they '• might readily be mistaken at first sight 
for distinct species " is erroneous. 

«, A. $ad., cC Ancla)nans(ir. Z>af/sow). A. O. Himie, Esq. [P.]. 
juv. sk8.; 

c. Iinm. sk. ; Koss I., Port Bliiir, S. R. G. Wardlaw Eaiii- 
skull. Andaiiuin. say [C.]. 

35. Pteropus niadiciis, MiUev. 

? Pteropus edulis {nee E. Geoff.), Nieuwenhuisen <^ lionenberff, J'erh. 

Batav. GenooUch. Kunst 4" Wet. xxx. p. 19 (1868 : Nias). 
? Pteropus [sp.], Rosenberg, Malay. Arch. p. 219 (1878 : Nias). 
Pteropus nieobaricus {nee Zelehor),Jentink, Cat. iSyst. Mamm. p. 147 

(pt.) (1888 : Nias) ; Modigliani, Ann. Mm. Civ. Geneva, (2) vii. 

p. 239 (1889: Nias); Trouessart, Cat. Mamm. i. p. 81 (pt.) 

(1897: Nias); Matschie, Megachir. p. 16 (pt.) (1899: Nias); 

Troues-fart, Cat. Mamm., Siippl. p. 50 (pt.) (1904 : Nias); Willink, 

Nat. rijd. Nederl. hid. Ixv. p. :375 (1905 : Nias). 
Pteropus niadicus, Miller , Proc. Biol. Soc. Wash. xix. p. C4 (1 May, 

1906: Nias). 

DUujnosis. — Similar to Pt. melanotus, but dentition rather heavier, 
back more or less aitproaching hair-brown, Hanks and sides of 
breast and belly pale sprinkled with blackish. Fore- 
arm 153-160 mm. Bab. Nias. 

Skull and teeth. — Skull not differing appreciably from that of 
Pt. melanotus. Teeth decidedly heavier (see measurements p. 231), 
but not differing in structure. 

Colour. — Three adult males, teeth much worn, skins (B. M.; U.S. Nat. Mus. 141231, '32, paratypes) :— Back and 
rump seal-brown, thickly mixed with greyish-white, mouse-greyish, 
or buffy-greyish hairs, producing the general effect of a shade more 
or approaching hair-brown. The amount of greyish admixture, 
though, so far as the material goes, always conspicuous, is subject 
to no small variation ; one sjiecimcu (141232) has the spinal tract 



sciil-brown, distinctly but not heavily sprinkled with greyish-white 
hairs, and (on tlie rump) M'ith tawny tips to the brown hairs, the 
admixture of paler hairs being thicker on the humerus and along 
tlie membranes ; in the Uritish Museum specimen the pale grej'ish 
are much in excess of the dark brown (in this specimen almost 
blackish) hairs, so as to make the whole of the back and rump 
appear light hair-brown or almost drab, sprinkled with blackish ; 
the third specimen (] 41231) is intermediate. — Middle of breast rich 
ochraceous, brightest (brilliant ochraceous-bufF or orange-buff) in 
the centre, gradually shading to tawny and from this to cinnamon- 
rufous or russet laterally. Anteriorly, towards the foretieck, the 
bright colour passes into chestnut more or less clouded with 
blackish, posteriorly, in anal region, into dull chestnut or chocolate 
more or less mixed with pale greyish. Extreme base of hairs in 
bright area seal- brown. Flanks and sides of breast and belly pale 
drab or drab-grey sprinkled with blackish hairs, or blackish heavily 
sj>riDklod witii pale drab or di'ab-grey. — Mantle chestnut more or 
less distinctly sprinkled with golden tawny hairs, shading poste- 
riorly, in a transverse line across shoulders, into tawny lightened 

External measurements 0/ Pteropus melanotus, tytleri, 
and niadicus. 

Ft. 'luelanotus. 

MiN. Max. 

I mm. 

Forearm ' 153 

Pillex, total length, c. u 65'5 

,, raetacra-pal | 14 

,, 1st pbaianx j 35 

2ncl digit, metacarjwl ' 80'5 

1st phalanx ,' \Qb 

„ 2nd-3rd phalanx, c. 11. .j 16 

3rd digit, metacarpal I 105'5 

,, 1st phalanx ; 77'5 

2nd pbaianx | 108 

4tli digit, metacarpal ! 101 

,, 1st phalanx : 61 

„ i'nd phalanx ! 63 

5th digit, metacarpal 109'5 

„ 1st phalanx , 45 

,, 2nd phalanx i 47"5 

Ear, length from oriiice ' (?27*) 

„ greatest width, flattened (? 19 *) 

Lower leg , 72 

Foot, c. u I 47"5 















Ft. tytleri. 
$ ad. 







Ft. niadicus. 

3 ad. I 

(Incl. two j 

paratypes.) , 

MiN. Max. 






























* From a slightly immature alcoholic specimen. 
t Estimate from skins. 



Jlectsuremcnts of skul 

and teeth of Pteropus melauotus, tytleri, 
and niadicus. 

Ft. metaiiotus. 
Skulls and 
teeth : 9 ad. 

Pt. tytleri. 
Skull : 1 ad. 
Teeth: 1 ad., 

1 iijim. 



Min. M.^i-K. 

Skull, total length to giiathion 
























3 8 






29 6 









mm. mm. 
(16 * 

,, palation to incisive foramina ... 
,, front of orbit to tip of nasals ... 
„ width of brain-case at zygomata. 
,, zygoaiatic width 


width across m\ externally 

lachrymal width 

width across canines, externally. 


9 5 

,, width of mesopterygoid fossa ... 

„ between p''-p', internally 

,, between cingula of canines 




Mandible, length 

„ coronoid height 


Lower teeth, c-ni., 


f pper incisors, combined width 

p', length 

„ width 

p^, length 

,. width 

ni", length 

,. width 

4-7 4-8 
3-5 3-8 
4-9 5 
3-7 3-9 
5-8 5-8 
33 36 

m-, leneth 

2-4 2-8 

„ width . 
p„ length . 
„ wiihh . 
p,, length 
I „ width 
1 Pi. lenglli 
;, width 
iDi, length 
I „ width 
'"o, length 
„ width 
m,, length 
„ width 





































Pt. niadwus. 

Skulls: 3 ad. 

Teeth : 2 ad. 

(Incl. two 


Min. Max. 









































232 rxEROPcs modiglianii. 

with orauge-buff. Sides of neck dark chestnut. Foreneck similar, 
hilt more or less strongly blotched with blackish. — Occiput similar 
to mantle (chestnut), shading into a rather brighter tinge (chestnut- 
russet) on crown. Muzzle and sides of head grizzled blackish and 
pale drab, in some specimens washed with russet in temporal 
region. Chin and throat blackish, with or without pale greyish 

Measurements. On pj). 230, 231. 

Specimens examined. Three, in the collections of the U.S. National 
(two paratypes) and British Museums. 

Range. The island of Nias, off W. Sumatra. 

Type in the U.S. National Museum (no. 141233). 

RemarJcs. — Undoubtedly a Nias representative of Pf. melanotus 
(Nicobar Islands). In Pt. melanotus a slight sprinkling of the 
blackish colour of the back, sides of breast and belly, and flanks 
with pale greyish is detectable ; in Pt. niadicus this admixture of 
pale colour has become so conspicuous as to change the general 
colour of the back into, or in the direction of, hair-brown, and that 
of the flanks and sides of breast and belly into pale drab. In 
Pt. melanotus the head, as far as back of ears, is usually blackish, 
but specimens occur in which the bright colour of the mantle 
extends forward on the crown, even to the front of the eyes; this 
latter has apparently become a fixed character in Pt. niadicus. The 
only other appreciable modification is the rather heavier dentition. 

a. S id. sk. ; Lelemboli, Nias; Aug. 1886 Marquis G. Doria 
skull. {Dr. E. Modigliani). [P.]. 

'- 36, Pteropus modiglianii, Thos. 

Pteropus modiglianii, Thotnaf, Ann. Mux. Civ. Gejiova, (2) siv. 

p. 105 (10 April, 1894 : Kifa-juc, Biia-Biia, Engano) ; Trouessart, 

Cat. Mamm. i. p. 81 (1807 : Engano) ; Miller, Proc. U.S. Nat. 

Mus. XXX. p. 8:23 (1906: Pulo Dua, Engano; habits); id., Fam. 

Si- Gen. Bati, p. 56 footnote, p. o8 (1907). 
Pteropus nicobaricus a. modiglianii, Matschie, Megachir. p. 17 (1899 : 

Engano) ; Trouessart, Cat. Mamm., Svppl. p. 50 (1904 : Eugano) ; 

Wi'llink, Nat. Tijd. Nederl. Ind. Ixv. p. 275 (1905 : Engano). 

Diagnosis. — Similar to Pt. melanotus, but fur longer, colour 
practically uniform blackish above and beneath, size smaller. Fore- 
arm 134"5-141 ram. Hab. Engano. 

Skull and teeth. — General characters of skull as in Pt. melanotiis, 
but size considerably smaller ; total length 61"5-67 mm., against 
72-7 5-5 in melanotus; actual size of orbits very nearly as in 
niadicus and melanotus, relative size therefore (considering smaller 
dimensions of skull) somewhat larger; diameter of orbit contained 
4'75-5"0, in niadicus and melanotus 5'4-5'55, times in total length 
of skull.— Teeth not reduced to the same degree as skull, therefore 
relatively larger than in the allied species (compare measurements, 
p. 236) ; structure of teeth quite as in melanotus and niadicus. 

Fi'.r. — I.ouger and more silky than in Pi. melanotus. Directed 
backward, but not closely adpret^^edj on back, spreading on mantle 


and rump. Longest hairs of back approximately 18, mantle 20, 
belly 20 mm. Distribution of fur as in Pt. melanotus. 

Colour. — Whole series examined (adult males and females, 
skins) : — General colour uniform blackish above and beneath ; 
mantle and middle of breast and belly not differing from rest of 
pelage. Sides of rump in most specimens more or less distinctly 
washed with dull vandyck-brown or mars-brown (in one specimen 
with ochraceous), this tinge occasionally extending forward on 
sides of back along membranes, and sometimes detectable also in 
the colour of the glandular hairs on 1 ho sides of the neck. 

Measurements. On pp. 235, 236. 

Specimens examined. Seven, from the collections of the U.S. 
National * and Bri1;ish Jiluseums, including one cotype of species. 

Range. Engano (and Pulo Dua, off S.E. Engano). 

Coti/pes in the Genoa (one) and British Museums (one). 

Ilahits. — In November 1904 Dr. W. L. Abbott found this species 
common in Engano, less common in the small islet of Pulo Dua, 
about one mile south-east of Engano. It was feeding on wild fruit, 
but did not frequent the cocoa-nut trees. 

a. 2 ad. al. ; skull. Bua-Bua, Engano; Marquis G. Doria [P.]. 
June, 1891 (jDi-. li. {Cotype of species.) 


37. Pteropus natalis, Thos. 

Pteropus natalis, Thomas, P. Z. S. 1887, p. 511 f, pi. xli. (animal) 
(Christmas I.) ; Lister, Nature, xxxvii. p. 20.3 (1887 : Christmas 
I.) ; id., P. Z S. 1888, p. 616 (Christmas I.); Thomas, torn. cit. 
p. 532 (Chi'istmas I.) ; Ridley, J. Straits Brayich P. As. Soc. 
no. 23, pp. 128, 130 (1891 : Christmas I.; habits); Trouessart, 
Cat. Ma?nm. i. p. 81 (1897 : Christmas I.) ; Andreics, Mon. 
Christmas 1. p. 23, fig. 7 (skull), pi. i. (animal) (1900: habits) ; 
Miller, Fam. 4' Gen. Bats, p. 58 (1907). 

Pteropus (Spectrum) hypomelanus I. natalis, Matschie, Megackir. 
p. 26 (1899). i. natalis, Trouessart, Cat. Mamm., Stipjil. p. 52 
(1904 : Christma.s I.). 

Large P'riiit Bat, Andrews, Geogr. Journ. xiii. p. 32 (1899 : Christ- 
mas I.). 

Diagnosis. — Allied to Pt. morlic/lianii, but smaller, M'ith shorter 
cheek-teeth and much longer fur. Blackish or seal-brown above 
and beneath, with or without a slight paler wash on the mantle. 
Forearm 125-135 mm. Hah. Christmas I. (Indian Ocean). 

SJcull. — General characters of skull as in Pt. niodiglianii, but 
size much smaller : total length 54-5-5G mm., against 61*5-67 in 
Pt. niodiglianii ; size of orbits (diameter contained 5'l-5"25 times 
in total length of skull) slightly larger than in Pt. niadicus and 
melanotus (5'4-5*55 times), but not (juite as large as in Pt. modi- 
f/lianii (4-75-5-0 times). 

Teeth. — Differing from those of Pt. niodiglianii in the following 
particulars : — Canines heavier at base, and more recurved ; cingulum 

• U.S. N.M. no?. HWTS, 80, 81, 80. 81, 85. 
t MiFprinteil Ft. mfh.s on p. 512. 

234 rxERorus xatalis. 

broader ; vertical groove on front face of upper cauines in most 
specimens quite obliterated, occasionally tracealtle on close exami- 
nation ; p, coinparativel)' larger, in" and m^ reduced, p'-m^ and 
p^-ra^ comparatively shorter, owing to a reduction of the posterior 
portion of the teeth ; iiosterior basal ledges of p" and p., distinct, 
though much shortened, in all other cheek-teeth scarcely developed. 
— All cheek-teeth, particularly the molars, in adult specimens 
covered with a dark brown or blackish coat of tartaric acid (some- 
times so thicl; as to nearly obliterate the structure of the teeth). 

Far. — Much longer than in Ft. modkillanii, dense, soft ; directed 
posteriorly on back, but not adpressed, spreading on mantle and 
rump. Longest liairs of back and mantle 25 mm., of belly 24 mm. 
Distribution of fur as in Ft. modiglianii, but hairs on upper surface 
of proximal third of forearm slightly more conspicuous, and fur on 
lateral interfemoral extending backward along inner side of tibia 
to end of proximal third or half of the limb. 

Colour. — Twelve skins, adult males and females (Feb., Aug., Sept., 
Oct.), teeth in different stages of wear: — General impression, imi- 
form blackish seal-brown above and beneath. Underparts (fore- 
neck, breast, belly, flanks) in some specimens precisely similar in 
tinge to back, in others distinctly paler, a dark shade of vandyck- 
brown approaching seal-brown. Mantle generally not differing 
from back and head, but in some specimens the tippet is indicated 
by a distinct russet or pale cinnamon wash of the hairs on the sides 
of the neck (above the neck -glands), and in some of these specimens 
this same tinge extends to tlie base of the fur of the whole of the 
mantle, while in others the paler tinge is confined to the sides of 
the mantle, the hairs in the central portion of the mantle being 
dark-coloured from tip to base. Eump generally not differing 
from back, sometimes faintly tinged with vandyck-brown. Fur 
everywhere very thinly sprinkled with greyish-white or greyish- 
buffy hairs, particularly on the underside of the bod)-. 

There is no appreciable sexual difference in colour. Three 
females (skins, teeth unworn or much worn ; Oct., Nov., one 
undated) have no trace of a tippet ; of nine adult males, five 
(skins, teeth unworn ; Feb., Aug., Sept., Oct.) have an indication 
of a tippet as described above, while four (same age ; Aug., Oct., 
Xov.) are similar to the females. 

A half-grown young male (skin) is similar in colour to the 
adults, and with no trace of pale colour in the mantle. 

Measurements. On pp. 235, 236. 

Specimens examined. The British Museum series. 
'_ Range. Christmas Island, S. of Java. 
', Type in collection. 

Habits. — This species (Dr. Andrews writes, 1900, 1. s. c.) is very 
common all over Christmas Island, hundreds of individuals being 
sometimes seen together. Near the settlement it causes consider- 
able destruction of fruit (papaias, bananas), but when the wild 
fruits, particularly those of the "Saoh"' { and the 
" Gatet" (Inocarpus) are ripe, comparatively few of these bats visit 
the gardens, while great numbers may be seen in the forest. They 



arc larRoly diurnul in their habits ; sevcrnl may be observo.l saili,,-. 

ihev'i;; ;'f 'f ' '^ ''f ^^ ^^^ ^^^ ^^^^ -..light: even at^id av aif 
. ;• 1 '-i^o f-eq«ently seen feeding in the daytime. The cry i^^ 
a very ioud harsh screech, apparently uttered both durin ' the-in 
.pirat.on as well as the expiration of the breath. Foetal specimen 
some near the term, were obtained towards the end of De eX a 
female carrying a well-gro^vn young one was shot at the end^f July 

$ arl. al. 

luill. al. 

'/. d" ad. sli. ; 

^'^;^::;^.l-«?.,:^ Lo,^sof,heAd,„iralty 

f,f- 6 ad., 
ad. ai. 

iiig Fish': Capf 

Christmas I. (ILM.S, 
' Flying Fish ' 
Capt. Maclear). 


ria '), 


(Type of species.) 

Loi-dsof the Admindtv 
[P.]. ^ 

87.4.2G.1' 3. 

J. J. Lister, Esq. [C. & 

S. ' Egeria '). 
<J-l.-6 6nA\ Clmstmas I.; Oct. Sir John Murray IP 1 
djuv.,25 1897, Feb. 1898 - '- -■ 

ad. ska. ; (Br. C. W. An- 

skulls of </--/.•. di-ews). 

'"■?..!""- ''"ST",,'' '"'• ^-"^'-"■'«"",m 

skulls. t^Br. Andrews). 

External measurements of Pteropus modiglianii and natalis., 

Pt. vwdigliaiiii. 

7 ad. 
(Incl. cotjpe.) 

Pt. natniis. 

12 ad. 
(Incl. type.) 




Pollex, total lengtii, c.uT ........ 

,, metacarpal 

)■ 1st phalanx 

2nd digit, metacarpal 

i> 1st phalanx 

! .. , ••. . 2nd-3rd phalanx, c. u , 1^0 

ord digit, metacarpal I 93.5 

.. 1st phalanx .......1 69-5 

2nd phalanx ' 99 

. 4th digit, metacarpal \'^_'^'^ g^^ 

.. 1st phalanx r..-; 

O 1^ , . I ^o 

^ J. Znd phalanx g^ 

oth digit, metacarpal ....."..!| 96-5 

» 1st phalanx I 40-5 

; „ 2nd phalanx 1 39 

I Ear, lengtii from ori fice ." ' .' 07 

I „ greatest width, flattened | 17 

Front of eye to tip of muzzle . ' 

Jjower Xee .... 



er leg 































92 5 











































Measurements of skulls and teeth of Pteropus modigliaiiii 
and iiatalis. 

Pt. modialianii. 
Skulls :" 7 ad. 
Teeth : 6 ad. 
(Iiicl. cotype.) 



Skull, total length to gnatbion 61-7 

palation to incisive foramina .. j 31 5 


front of orbit to tip of nasals 
width of brain-case at zygomata 

zygomatic width ! 

width across m\ externally 

lachrymal width 

width across canines, externally j 12-6 

postorbital constriction 6'5 

interorbiral constriction | 8-2 

width of niesopterygoid fossa ! 7 

between p''-p-i, internally j 10 

,, between cingula of canines 

„ orbital diameter 

Mandible, length 

,, coronoid height 

Upper teeth, c-m^ 

Lower teeth, c-nig 

Upper incisors, combined width 
p', length 

,, width 

p'', length 

„ width ..'' 

m' , length 

„ width 

m^, length 

„ width 

Pi, length 

„ width 

Pa. length 

,, width 

p^, length 

,, width 

mj, length 

„ width 

ffij, length 

„ width 

fflj, length 

„ width 









66 8 


















5 3 


Pt. vatalis. 

Skulls and 

teelh : 12 ad. 

(Incl. type.) 











4-8 » 





































■* Senile sjiecimcn. 


F. Thk Ptkhopvs melasopogon group. 

Species. — Pt. melanopofjon, aruensis, keyensis, livingstonei. 

Range. — Amboina group (perhaps north-west to Sanghir Islands), 
Banda Islands, Timor Laut, Key and Aru Islands. One species iu 
the Malagasy region (Comoros). 

General characters. — Skull typical Pteropine (rostrum somewhat 
shortened in Ft. keyensis). dentition heavy but not otherwise modi- 
fied. Palate-ridges normal (54-5 + 3). Ears moderate, broader 
than usual, not attenuated above ; tip rather blunt, in one species 
extremely broadly rounded otf. Fur generally very short and 
closely adpressed ; tibia always naked above. Interfemoral scarceh' 
developed in centre. Colour varying according to species. Sexual 
differentiation inconspicuous (canines heavier in males, no neck- 
tufts). Size large or very large (forearm 162-204 mm.). 

Specific differentiation. — Pt. melannpiogon is peculiar in having 
the fur of the back exces.sively shortened and restricted to a narrow 
spinal tract (aged specimens in abraded pelage being nearh' naked 
on back); in the other species the furred area of the back is of 
normal breadth. In colour Pt. melanopogon is the most ordinary- 
looking of the four species : blackish back, chestnut head and 
collar, golden-buffy underside of body (compare Pt. rufus). Pt. aru- 
ensis is not essentially different from Pt. melanopogon in the colour 
of the head, collar and underparts, but the back is glossy silvery 
whitish (an excessive development of the silvery whitish element 
in the colour of the back is seen in certain other species of the 
genus : compare Pt. hypomelamis canus and lepidus with the dark- 
backed races of the same species, certain individuals of Pt. hypo- 
melamis tomesi with the normal dark-backed individuals of the 
same race ; even in Pt. a7-nensis the back is not uniform silvery 
"whitish, but thinly sprinkled with blackish hairs, reminiscent of 
the descent of the species from a dark-backed form). Pt, keyensis 
ia nearly uniform yellowish above and beneath. As already men- 
tioned above, the ears in the typical species of this group are rather 
bi'oader than usual with the tip somewhat blunt ; in the only 
Malagasy representative {Pt. livingstonei) they are so excessively 
broadened above as to be almost semicircularly rounded off, the fur is 
longer and harsher than usual, the general colour blackish above and 
beneath, with or without a remnant of a bright tippet; it is a 
mountain form, confined to the Comoros, perhaps to one island 
(Johanna I.). 

Affinities of group. — Judging from the typical Pteropine cha- 
racters of the skull and dentition, the origin of this group is pro- 
bably close to that of the foregoing types of the genus. It is not 
improbable that the rufus, melanotus. and melanopogon groups are 
rather intimately related (compare the style of colour of Pt. melano- 
pogon with that of Pt. rufus), the first representing a truly Malagasy, 
the second an Andaman-Nicobar, the third an Austro-Malayan branch 
of the same prototype. Pt. melanopogon, keyensis, and aruensis have 
hitherto been placed in the closest vicinity of Pt. 2W2^unnus and 

23S I'TEKoPu.s MEi.Axoroaox. 

neohiheniimtt {origmaWy these five species were described as varieties 
of one form): hntPt. pa/iuanus and neohihcrnicus are so essentially 
differeut from the three former species in dentition and palute- 
ridges (5 + 8-f 3) that a close relationship witli the melanopogon 
grou]) would seem to be excluded. 

38. Pteropus melanopogon, Pet. 
l'tero];)us melanopocjon (pt.)) Dobsou, Cat. Chir. B. M. p. 44. 

Pteropus pliaiops {nee Temm., 1825), Temminck, Mon. Maimn. ii. 
p. 65 (pt.), pi. XXXV. tig. 8 (head), ^\. xxxvi. figs. 1,2, 3 (skull) 
(1837 : Amboiua) ; Wagner, Schrehers Siiiiff., Siippl. i. p. 34(3 (pt.) 
U839 : Amboija) ; <S'. Aliiiler, in TommincJc s Nat. Gesch. Nederl. 
Ocerz. Bez., Zoorjd. pp. 20, 59 (pt.) (1839-44 : Amboina ; Banda) ; 
Lesson, N. Tab/. R. An., Mamm. p. 13, no. 171 (pt.) (1842: Am- 
boina) ; Schinz, Si/st. Verz. Siimj. i. p. 122 (pt.) (1844 : Amboina ; 
Banda): Gray, Zool. ' Samara7i(/,' Vert. p. 11 (pt.) (1849: Am- 
boina; iiandaj; Wagner, Schrebers Suiu/., Suppl. v. p. 596 (pt.) 
(1853-55: Amboina; Banda); Fitzinger, SB. Akad. Wien, Ix. 
Abth. i. p. 418 (pt.) ( 1870 : Amboina ; Banda) ; Rosenberg, Malay. 
ylrc/i. pp. 322, 360 (1878 : Ceram ; Goram). 
1 Pteropus phajops (pt.), Giebel, Sdug. p. 996 (1855: Amboina). 

Eunycteris pliaiops var., Gray, Cat. Monk. iJJc. p. 113 (1870: Buru ; 
Ceram; Goram). 

Pteropus melanopogon, Beter-i, MB. Akad. Berlin, 1867 (27 May), 
p. 330 (Buru; Amboina; I3oano ; Saparuf) ; Ceram; Goram; 
Manavolka); Dohson, Cat. Chir. B. 31. pp. 44,47 (pt.) (1878: 
liuru ; Ceram ; Goram ; the locality '• Ceram " lor .specimen d is a 
mistakfi for Goram) ; Truuessart, Rev. ^- Mag. Zool. (3) vi. p. 205 
(pt.) (1879: Amboina; Boano ; Saparua; Ceram; Goram; Mana- 
volka); Jcntink, Cat. Osf. Mamm. p. 255 (pt.) (1887: Siao, 
Sanghir Is. ; Burn ; Amboina ; Boano : Ceram ; Goram ; Mana- 
volka) ; id. Cat. Syst. Mamm. p. 142 (1888: Siao: Buru; 
Amboina; Boano; Saparua; Ceram; (ioram ; Manavolka); 
Troucssart, Cat. Mamm. i. p. 80 (pt.) (1897 : Buru; Amboina; 
Ceram) ; Willink, Nat. Tijd. Nederl. Ind. Ixv. p. 272 (pt.) (1905 : 
Buru ; Amboina ; Ceram ; Goram ; Banda) ; Miller, Bum. i)- Gen. 
Bats, p. 58 (1907). 

Pteropus (Eunvcteiis) melanopogon, Matschie, Megachir. p. 11 
(1899 : Burii ; Amboina) ; Trouessart, Cat. Mamm., Suppl, 
p. 49 (1904 : Sanghir; Buru; Amboina; Ceram). 

Diagnosis. — Skull typical Pteropine. Posterior ledges of pre- 
molars above and below well marked but short ; cingulum of 
canines well defined but rather narrow. Ears moderate, broadly 
rounded ofi" above ; wing-membranes arising very close together 
from sides of spine ; interfcmoral short in centre ; fur of back 
extremely short, adpressed, and restricted to a very narrow spinal 
line ; forearm and tibia practically naked. Back blackish ; head 
and neck all round rich chestnut lightened with golden orange-buff 
above ; breast and belly generally uniform light golden bufly, some- 
times buffy in centre and darker on sides ; chin and throat blackish. 
Size very "large: forearm 196-204 mm. Hah. Amboina group; 
Banda Is. ; Timor Lnut, ) j^jj :,, ;;., ,.',j ir-.-i ; 


Slidl. — Heavily built, with large orbits, strong crests, and long 
postorbital processes, llostrum deep and broad ; distance from tip 
of postorbital processes to gnathiou more than from former point to 
lambda. Orbital diameter greater than width of rostrum across 
alveolar borders of p'-p^ ; front of orbit above front of m^ Post- 
orbital processes nearly reaching, sometimes in contact with, 
corresponding processes of zj'goma ; frontal region between post- 
orbital processes much concave. Sagittal crest deep. Palate much 
broader than in Pt. vampyrus -. between inner sides of p^-p'' 
14-14'8 mm., against 12'2 to 12"8 in Pt. vampyrtis vampyrus. 
Coronoid process bioad antero-posteriorly, but not higher than usual ; 
coronoid height of mandible less than c-mj ; condyle considerably 
above alveolar line of mandible. 

Teeth. — Cingulum of upper canines well defined but rather 
narrow ; crown in profile slightly recurved ; median ridge on Hngual 
face and anterior and posterior margins sharply projecting ; vertical 
groove on front face broad and deep, terminating well above tip of 
tooth, p' spiculiform, deciduous. Posterior ledge of p^ and P3 
short, distinctly marked off postero-cxternally by small notch from 
base of outer maiu cusp. Corresponding ledge of p' and p^ shoit, 
more developed on inner than outer side of tooth ; postero-external 
notch generally obsolete, sometimes (at least in young teelh) quite 
distinct; line of posterior margin of p' projected inward passing 
very nearly through middle of m' of opposite side, m' nearly once 
and a half as long as broad, nr equal to or slightly larger than p^. 
i^ once and a half or nearly twice the bulk of i^. Lower canines 
recurved, cingulum well defined, but rather narrow. Pj from twice 
to nearly twice and a half the size of i,. m.^ generally a trifle 
smaller than Pj. 

Palate-ridges. — 5-|-5-f3. First ridge terminating laterally at 
front of canine ; second at back of canine ; third at front of p' ; 
fourth at back of p^ ; fifth at middle or front of p'; sixth at front 
of m' ; seventh at back of m' ; eighth to tenth behind m'- ; eleventh 
to thirteenth situated at palation border. 

Ears. — Moderate, exposed, reaching about half the distance to 
hinder corner of eyes. Upper third of inner margin strongly 
convex, corresponding portion of outer margin flatly convex, 
making tip of conch rather broadly rounded off. 

WiiKjs. — ilembranes arising unusually close together (about 
lU mm. apart) from sides of spinal region. 

Jnterftiiioral. — Much reduced in centre ; depth a few milli- 

-Pa?'. — Extremely short on upperside of body, and restricted to a 
narrow spinal line of closely adpressed hairs ; moderate on neck and 
underparts. Length: back 7-8, mantle 16-1 1), belly 13-17 mm. 
Least width of furred line of back !)-ll mm. Specimens in abraded 
pelage are nearly naked cm middle of back. 

Forearm, tibia, and tip of femur naked above and beneath, except 
for some thinly scattered, short, adpressed hairs on upperside of 
forearm in front of elbow region. Interfcmoral naked, except iu 

240 rxERorus mklaxopogo.v. 

centre and along inner side of femur. Hairing on tmdcrside of 
membranes as usual. 

Colour. — Skins of two adults, Burn and Ceram ( and : Back glossy blackish seal-brown ; rump slightly mixed 
with chestnut, inner sides of thighs with chestnut and greyish 
huffy hairs. — Breast rich glossy golden yellowish-huff, strongly 
contrasting with chestnut foreneck, shading gradually into glossy 
golden straw-yellow on belly and crissum, and into orange-ochra- 
ceous on flanks. Base of fur everywhere nearly straw-yellow. 
Underside of humerus blackish sprinkled with huffy. — Ifantle rich 
glossy chestnut mixed with golden orange-buff hairs, lightening 
])osteriorly (shoulder-region) to deep golden orange-buff, and passing 
on sides of neck into dark chestnut, this in turn on foreneck into 
a slightly brighter golden chestnut. Concealed bases of hairs 
everywhere paler than tips. — Crown glossy yellowish orange-buff 
(somewhat paler than shoulders), darkening on sides of head to 
chestnut sprinkled with orange-buff hairs, this again to blackish on 
throat and chin. 

A nearly new-born young from Burn (skin, ; hairs not 
out on centre of breast and throat) is similar in colour to the 
adults, hut with paler mantle and head. 

An immature female from Goram (skin, is similar to 
the adult specimens from Burn and Ceram, but with underside of 
body conspicuously darker. Centre of breast and belly rich glossy 
golden buff-yellow, gradually shading into tawny ochraceous and 
tawny on sides of breast and belly, and this tinge again into deep 
vandyck-brown on flanks and anal region. Underside of humerus 

An immature male from Timor Laut (al., is perfectly 
similar in colour to the above described adult specimens from Burn 
and Ceram. 

The type of the species (Amboina) and five specimens in the 
Leyden Museum from Amboina, Ceram, and Saparua are on the 
whole very similar to the British iluseum series from Burn, Ceram, 
Goram, and Timor Laut, the only noteworthy individual variations 
being nearly the same as in the British Museum series, viz., the 
lighter or darker tinge of the colour of the head, mantle, and flanks, 
and the more or less conspicuous sprinkling of the inner sides of 
the thighs with chestnut and buffy hairs. The colour of the flanks 
varies in the whole series from rich orange-ochraceous (brightest 
extreme) to nearly chestnut seal-brown. 

Seanud differeutiatlon. — Canines noticeably stronger in males 
than in females. Measurements taken on one male and tbree 
females : upper canines, vertical extent from alveolar border 
12 mm. (male) and 10-11 (females), antero-posterior basal diameter 
6 (male) and 5-2-5'6 (females) ; lower canines, vertical extent 10'5 
(male) and 8-9 (females), antero-posterior basal diameter 4*5 (male) 
and 3-8-4 (females). 

Measurements. On pp. 244, 245. 

^j.'£ciiiuiis exmidned. Thirteen, in the collection? of the Berlin 


(one, the type, Amboina), Leyden (seven, Aml3oina,Ccram, Sapania, 
viz. mounted specimens <j, h, i, j, I, skeletons a, h), and British 
Museums (five, Buru, Ceram, Gorara, Timor Laut). 

Range. Amboina group : Burn, Amboina, Ceram, Boano, Saparna, 
Coram, Manavollia ; Banda Islands ; Timor Laut. The species has 
been recorded in literature from the Sanghir Islands ; see references, 

TyjJ'i in the Berlin Museum. 

Pteropvs n)eIano2Jogon, Peters; 1867. — Type locality, Amboina. 
Tj'pe, a mounted young adult male, skull in situ, collected by 
S. Miiller, and acquired from the Leyden Museum ; Beg. no. 2627. 

Remarks. — The broad rostrum, AvcU-markcd posterior ledges of 
the premolars, broad ears, excessively narrow furred area of the 
back, and very large size, are the most conspicuous diagnostic 
characters of Ft. melanopor/on. In the reduction of the fur of the 
back it is approached by Pt. papuanus and neohibeniims, in which, 
however, the posterior ledges of the premolars are obsolete ; in 
size it is surpassed only by the largest races of Pt. vampyriis. 
The species has rather closely allied representatives in the Aru 
{^Pt. aruensis) and Key Islands (Pi. Iceyensis). 

Purcbasetl (Baker). 

Dr. A. E. Wallace [0.]., 3. 

Purchased (Verreaux). 
Dr. A. E. Wallace [C.]. 

British Association [P.]- 

39. Pteropus aruensis, Pd. 
Pteropus melanopogon (pt.), Dobson, Cat. Chir. B. M. p. 44. 

Pteropua argentatus (nee Gray, 1844), Gray, P. Z. S. 1858, p. 107 
(Aru Is.) ; id., Cat. Mamm. New Guinea, p. 2 (1859 : Aru Is.) ; 
Finsch, Neu- Guinea, p. 150 (pt.) (1865: Aru Is.) ; Rosenberg, 
Reis. Zuidoostereil. p. 31 (pt.) (1867 : Aru Is.) ; id., Malay. Arch. 
p. 360 (pt.) (1878: Aru Is.). 

Pteropus melanopogou var. aruensis, Peters, MB, Akad, Berlin, 
1867 (27 May), p. 380 (Aru Is.) ; Peters S) Doria, Ann. Mus. Civ. 
Genora, xvi. p. 689(1881 : Aru Is.) ; Troiicssarf, Cat. Mamm. i. 
p. 80 (1897: Aru Is.); Matschie, Meyachir. pi. vi. figs. 3, Za, 
3 6 (skull) (1899). 

Pteropus aruensis, Jentink, Cat. Ost. Manim. p. 256 (pt.) (1887: 
Wokam ; Wonumbay) ; id.. Cat. Syst. Mamm. p. 143 (pt.) (1888 : 
Woliam ; Wonumbay); Matschie, Mf.gachir. p. 14 (1899: Aru 
Is.); Tronessart, Cat. Mamm., Suppl. p. 49 1 1904: W^okam) ; 
Willinh, Nat. Tijd. Nederl. Ind. l.vv. p. 274 (1905 : Aru Is.). 

Pteropus melanopogon aruensis (pt.), Heller, Abh. Mus. Dresden, vi. 
no. 8, p. 4 (1897 : Aru Is.). 

Pteropus rubiginosus, Rosenberg, Reis. Zuidoostereil. p. 31 (1867: 
Wokam) ; id., Malay. Arch. p. 360 (1878 : Wokam). 

a. [$] ad. skull. 

b, c. $ ad., pull. 



d. Ad. St. 


€. 5 imni. sk. ; 



f. (5' imm. sk. in 

Timor Laut (Dr. 

al. ; skull. 

H. 0. Forbes). 

242 rrj'.ROPi'f; akvexsis. 

P Pteropus funugiiUis, Boseiiberf/, liein. Zuu/uostcreil. p. ol ( 18(37 : 

Wonumbay) ; id., Malay. Arch. p. 3(30 (1878 : Woiiumbay). 
rteropus hypomelas {nee Pt. hypomelanus, Tvmm.),Graij,Cat. Monk. 

i^-c. p. 110 (1870: Aml8.). 
Pteropus melanopogon vav. /3 (pt.), Dobsoii, Cat. CJiir. B. M. pp. 4o, 

47 (1878 : Aru Is.). 
Pteropus nielauopogon (pt ), Troiiessart, Rev. 8,- Mag. Zool. (3") vi. 

p. '20o (1879 : Aru Is.) ; Willink, Nat. Tijd. Nederl. Ind. Ixv. 

p. 272 (1905 : Aru Is.).' 

Diagnosis. — Similar to Pt. melanopogon, but furred area of back 
not Tinxisually narrow, colour different. Back and rump glossy 
silvery whitish, contrasting Avith chestnut collar. Forearm about 
190-191 mm. Hah. Avu Is. 

Skvll and teeth. — Apparently not differing in any noteworthy 
character from those of Pt. melanoporion. The three skulls examined 
(Leydeu Museum, n, h, c) are slightly immature. 

Pur. — Closely adpressed on back, but longer thau in Pt. melano- 
■poqon. Approximate length, back 12-13, mantle and belly 16- 
17 mm. I,east width of furred area of back 45-47 mm. Distri- 
bution of fur on limbs as in Pt. melanopogon. 

Colour (type). — Back and rump glossy silvery greyish-white, 
everywhere thinly sprinkled with blackish hairs [-'back of a silky or 
silvery shining wbite, very beautiful in the freshly killed animals," 
AVallaco MS. quoted by Gray, I. s. c.\ — Flanks and belly golden bufFy 
slightly washed with pale cinnamon, gradually darkening to cinna- 
mon-rufous on lateral parts of front of breast, and clouded with 
blackish chestnut on centre of breast. Breast, belly, and flanks 
everywhere sprinkled with glossy pale golden buffy and a few 
blackish hairs.- — Mantle blackish maroon-chestnut, lightening to 
chestnut posteriorly in shoulder region and anteriorly on occiput, 
and gradually darkening to blackish seal-brown on sides of neck 
and foreneck. Base of fur of collar everywhere much lighter than 
exposed tips of hairs. — Crown and sides of head dark brownish, 
heavily sprinkled with shiny pale buify hairs. Chiu blackish seal- 
brown like foreneck. 

Individual variation chiefly confined fco a darker or somewhat 
paler tinge of the chestnut-coloured pDrtions of the pelage. Back 
and rump silvery whitish in all specimens examined. 
Measurements. On pp. 244, 245. 

Specimens examined. Six, in the collections of the Berlin (two), 
Leyden (three), and British Museums, including the type of the 

Range. Aru Islands (Wokam). 
Type in the Berlin Museum. 

Pteropus argentatus, Gray; 1858. — Pteropus argentatus, Gray, 
1844, is a species allied to Pt. dohsoni and Pt. caviceps (sec 
antea. p. 197). Pt. argentatus, Gray, 1858 (I. s. c), is the species here 
under consideration, as jjroved by his description and the specimen 
he had before him and which is in the collection. There can be no 
doubt that Gi'ny, in 1858, correctly considered this bat an undescribed 

PTKUOPUs AUi-i;xsis. 243 

species, ami that it was only by a slip of the memory that he gave 
it a name which lie had, fourteen years earlier, proposed for a 
different species of the geiuis. In 1870 (Cat. Monk. &c. p. 110, 
under the synonyms of Pl.hi/pomelas) he had discovered the lapsus, 
but now committed the error to identify the bat with the 
widely different Ft. ht/poindaniis, Temm., the technical name of 
which he wrote Pt. hypomehis. 

Plerojjiis r.i l't)i<>po[/on var. aniejisls, Peters ; 1SG7. — -Type : J ad, 
mounted, skull in situ, Am Islands, collected by Bernstein, acquired 
by the Berlin from the Leyden Museum, lleg. no. 4962. The 
figure of the skull iu the ' Megachiroptcra des Berliner Museums ' 
{I. s. c.) is not drawn from the type, nor from the only other 
specimen in the Berlin Museum (4703, Aru, Dr. Beccari) ; the 
original may be a Leyden skull (?). — Dobson (Cat. Chir. 1878) 
followed Peters, iu so far as he considered Pt.aruensis only a variety 
of Pi. melanopo(ion, but with this variety he united Pt. keyensis. 
Jentink's view (Cat. Ost., 1887; Cat. Syst., 1888) is only a slight 
modification of Dobson's ; he recognized Pt. aniensis as specifi- 
cally distinct from Pt. melanopogon, but uuited with the former 
Pi. kei/eiisv;. Matschie (1899) recognized P<. nielanopojon, aruoisis, 
and ke)jen.sis as distinct species, but the former he placed in the 
subgenus Eimi/iUeris, the two latter in the subgenus Pteropus. 

Pterojius rubirfinosus, Hoscnherg; 1867. — Described by Rosenberg 
as uniform " roestkleurig," darkest on head and neck, palest on 
underparts, iris brownish red ; somewhat larger than '' Pt. argen- 
toYies" [which in Rosenberg's writings is synonymous withPi. aruensis 
and Pt. heijensis, as shown by a series of specimens from the Aru 
and Key Islands in the Leyden Museum, collected by Rosenberg, 
and labelled Pt. ar(jentatiis\ One example only, Wokam, Aru 
Islands ; original number 23. In Rosenberg's manuscript Catalogue 
preserved in the Leyden Museum this specimen (Wokam, no. 23) is 
described as " roestgeel," larger t ban Pt. argentatiis, iris brownish 
red, 9th April, perhaps a new species. The type is not in the 
Leyden Museum. By Matschie {I. s. c.) referred, with a query, to 
Pt. aruensis, but Rosenberg's brief notes (as quoted above) would 
rather seem to indicate a species allied to Pt.jJapnanus. — Rosenberg's 
' Reis naar de Zuidoostereilanden ' (dated 1867) appears to have been 
published late in 1867 or early in 1868, in either ease after the May 
issue of the ' Monatsberichte ' of the Berlin Academy for 1867 which 
contained Petcrs's description of Pt. melanoj^ogon var. artansis. 

Pteropus fiimigatus, Rosenbei'g ; 1867. — Described by Rosenberg 
as " graauw-bruin " (in the German version, 1878, " einfarbig 
rauchbraun ''), smaller than Pt. argentatu', iris brownish yellow; 
common in Wonumbaj , Aru Islands. In Rosenberg's MS. Catalogue 
of his collections (Leyden Museum) only one specimen is registered : 
Wonumbay, no. ;')6, female, " grijsachtig bruin," below medium size, 
iiis yellow, 15th May. The type is not in the Leyden 
Museum. By Matschie {l.s.c.) put down, with a query, under 
Pt. aruensis, a suggestion certainly not supported by Rosenberg's 
ijotp on the colour and size. Perhai)s a distinct species. 




JZemarlcs. — The collar (mantle, sides of neck, and foreneck) is in 
this species rather similar in colour to that of Pt. melanofogon, 
though on the whole slightly darker. But the chestnut of the collar 
is made still more conspicuous by the contrast of the beautiful silky 
whitish tinge of the back, a colour rather rare in the genus (cf. Pt. 
temmincki, capisiraUis, personafus, hypomelanus canus, h. lejndus). 
There can be no doubt that Pi. aruensis is the Aru representative 
of Pt. melaiiopor/071, with which species it completely agrees in the 
characters of the skull and teeth and the shape and size of the ears. 
Pt. aruensis is similar to many other species of the genus in the 
distribution of the fur of the back, but peculiar in colour, while 
Pt. melanojjor/on is rather ordinary in colour, but peculiar in the 
extreme reduction of the furred area of the back. 

a. Ad. St. 

Aru Is. 

Dr. A. R. Wallace [0.]. 

{Tj/pe oi' Pf. argentatus, Gray, 1858 nee 1844.) 

External measurements of Pteropus melanopogon and aruensis. 

Pt. melanopogon. 
3 ad.* 

Pt. aruensis. 



inni. mm. 

Forearm ' 196 204 

PoUex, total length, c. u 1 74-5 81 

metacarpal [ 17-5 20 

„ 1st phalanx • 405 41-5 

2nd digit, metacarpal | 103 108 

iBt phalanx i 21'5 24 

2nd-3rd phalanx, c.u...! 19 20 

3rd digit, metacarpal 134 144-5 

l8t phalanx : 97-5 104-5 

2nd plialanx j 144-5 147 

4th digit, rn etaca rpal i 1 28-5 136 

Ist phalanx \ 80-5 86-5 

,, 2nd phalanx I 81-5 85 

5th digit, metacarpal i 137 147 

,, Istphalau.x ! Gl (13-5 

2nd phalanx I CO 63-5 

Ears, length from orifice I 30 1 

,, greatest width, flattened ... 19-6 1 

Lower leg 835 85 

Foot, c.u ... 68 

Calcar ... 24 






























132 5 





* B.M. (Burn) and (Cerara); Leyden Museum / (Amboina). 
t Type of species, and B.M. 
J Estimate (skins). 



Measurements of skulls of Pteropus melanopogou, and teeth of 
Pt. melanopogon and aruensis. 

Skull, total leuglh to gnatbion 

„ palation to incisive foramina ... 
„ front of orbit to tip of nasals ... 
,, width of brain-case at zygomata 

„ zygomatic width 

,, width across m^, externally 

„ lachrymal width 

,, width across canines, extemaliy ... 

„ postorbital constriction 

,, interorbital constriction 

,, width of mosopterygoid fossa .. 

„ between p^-p^, internally 

,, between cingula of canines 

„ orbital diameter 

Mandible, length 

„ coronoid height 

Upper teeth, c-m- 

Lower teeth, c-mj 

p3, length 

„ width 

pS length 

„ width 

m', length 

,, width 

111^, length 

„ width 

Pi, length 

„ width 

Pj, length 

,, width 

P4, length 

„ width 

iDj, length 

„ width 

m^, length 

,, width 

HI J, length 

,, width 

P(. melanopogon. 

Skulls: 2 ad. 

Teeth: 2 ad,, 

2 iium.* 






5 8 


2 2 






2 5 

Pt. aruensis. 
Skulls : none. 
Teeth : 3 imm. 

MiN. Max. 





















6 7 



* 2 ad., viz. B.M. (Burn), Leyden Museum skeleton a (Amboina) ; 
2 imm., viz. B.M. (Gorara) and (Timor Laut). 
t Leyden Museum, teeth of skulls a, b, c. 

246 FiEEorirs kuyessis, 

40. Pteropus keyensis, Pet. 
Pieropns melanopogon (pt.), Dobson, Cat. Chir. B. M. p. 44. 

Pteropus melanopogon var. keyensis, Peters, MB. Akad. Berlin, 

1867 (27 May), p. 330 (Key Is.) ; Peters <^ Boria, Ann. Mm. Civ. 

Genova, xvi. p. 689 (1881 : Kev Is.) ; Tronessurt, Cat. Mamvi. i. 

p. 80 (1897 : Key Is.). 
Pteropus keyensis, Mutschie, Megachir. p. 14 (1899: Key Is.); 

Trouessccrt, Cat. Mamin., Suppl. p. 49 (1904 : Great Key ; Little 

Key : Koor) ; Willink, Nat. Tijd. Nederl. Ind. Ixv. p. 274 

(1905 : Key Is. ; Koor). 
Ptercpiis avgentatiis (pt., nee Gray), Posenberg, Peis. Zvidoostereil. 

p. 31 (1867: Key Is.); id., Malay. Arch. p. 300 (1878: 

Key Is.). 
Pteropus melanopogon vnr. 3 (pt.), Dobson, Cat. Chir. B. M. p. 45 
; (1878: Key Is.). 

Pteropus melanopogon (pt.), Trouessart, Rev. ^- May. Zool. (3) vi. 

p. 205 (1879: Kev Is.) ; Willink, Nat. Tijd. Nederl. Ind. Ixv. 

p. 272 (1905 : Key Is.). 
Pteropus aruensis (pt., 7iec Pet.), Jentink, Cat. Ost. Mamm. p. 250 

(1887 : Great Key : Little Key ; Koor) ; id., Cut. Syst. Mamm. 

p. 143 (1888 : Great Key : Little Key ; Koor). 
Pteropus melanopogon aruensis (pt.), Heller, Abh. Mus. Dresden, vi. 

no. 8, p. 4 (1897: Key Is.). 
I Pteropus elirysargyrus, Ileude, Mem. Hist. Nut. Pmp. Chin. iii. 

p. 177, footnote, pi. v. fig. G (teetb) (1896 : Little Key). 

Diar/nosis. — Similar to Pt. a7'uensis, but colour of fur nearly 
uniform light yellowish above and beneath. Forearm 179-187'5 mm. 
Ilab. Key Is. 

SkuU and teeth. — Rostrum markedly shorter than in Pt. melano- 
pogon: from front of orbit to tip of nasals 25-27-7 mm., against 
28"2-31'2 in Pt. melanopogon; front of orbit vertically above back 
of p\ in Pt. melanopogon above front of m'. Teeth not differing 
appreciably from those of Pi*, melanopogon and aruensis. 

Palate-ridges. — As in Pt. melanopogon. 

Wings. — Membranes less closely approximated than in Pt. melano- 
pogon, about 21 mm. apart at origin from sides of back. 

Fur. — Length and distribution of fur as in Pt. ainunsis. Approxi- 
mate length, back 11-13, mantle and belly 14-15 mm. Least 
■width of furred space of back about 45-47 mm. 

Colour. — c5' ad. al., teeth worn, Back and rump yel- 
lowish cream, tinged with pale maize-yellow on back and with buff- 
yellow on rump. — Fur of breast, belly, flanks, and crissum cream- 
coloured at base, tipped with buif-yellow on breast, belly, and flanks, 
and with light straw-yellow on crissum. Underside of humerus and 
woolly hairs on underside of membranes pale buify. — Mantle, sides 
of neck, and Ibreneck deep chrome, palest on foreneek, shading into 
light buft'-ycllow on crown and sides of head, and into cream on 
muzzle and throat. 

Individual variation in colour apparently small. All the indi- 
viduals examined differ only in the deeper or paler tinge of the 

PTERorrs LivixGsroxEi. 94.7 

Menswements. On pp. 248, 241). 

Specimens examined. Sixteen, in tlie ccllcctions of the EerJin 
(three), Leyden (twelve), and liritisli Museums, including the type 
of the species. 

Hetnge. Key Islands : Great Key, Key Doulan, Little Key, Koor. 

Tyjjfi in the Berlin Museum. 

Fteropus melnnopogon var. Jcetfeusis, Peters ; 1867. — Ty])e locality, 
Grand Kej'. Type, d' ad., mounted, skull extracted, collected by 
Rosenberg, acquired from the Leyden Museum ; llcg. no. 4752. 

Fteropus chri/sargi/rus, Heude ; 189G. — Type locality, Little Key. 
Cotype-s (at least two) presumably in the Zi-ka-wei Museum, 
Shanghai. The description of colour and size, and figure of teeth 
agree perfectly with Pt. l-ei/eDsis. 

liemarls. — This species is the Key Island representative of the 
Ft. melanopor/on type. It accords with Ft. mclanojjogon and ciruemns 
in the characters of the skull (apart from some shortening of the 
rostrum) and teeth, the number and arrangement of the palate- 
ridges, and the size and shape of the ears. It accords with Ft. aru- 
cnsis and diifers from Ft. melanopoi/on in the less closely approxi- 
mated wing-membranes and broader furred area of the back. And 
it differs from both of the related species in the highly peculiar pale 
yellowish colour of the fur. 

rt. (5 ad. al. ; Kev Doiilan, Key Is. Lords of the Treasury [r.l. 82 7.27 3. 
.skull. (H.M. . • Challenger '). 

41. Pteropus livingstonei, Crrai/. 
Fteropus llvivgstonii, Dobson, Cat. Chir. ]>. M. p. 55. 

Pteropus livlngstouii, Grny, P. Z. S. 186G, p. G6 (Johanna I.) ; 

id., Cat. Monk. Sfc. p. 109 (1870: Johanna); I)obso?i, I. s. r. 

pi. iii. fig. 6 (ear) (1878 : Johanna) ; Trunessart, Pev. Sf Maej. Zool. 

(3) vi. p. 202 (1879 : Johanna) ; Bobson, Pep. Prif. Assoc. 187», 

p. 162 (1879 : remarks on distribution); Jentink,Cat. Si/sf. Mamm. 

p. 147 (1888: Johanna): Mibie-Edivarch ^- Oustal'et, N. Arch. 

Mus. d'Hist. Xat. Paris, (;2 ) x. p. 224 (1888 : Johanna) ; Tnmessart, 

Cat. Mamm. i. p. 81 (1897: Johanna); Matschie, Miu/arhir. 

pi. iv. figs. 9, 10 (skull) (1899: Johanna); Miller, Fam.\ Gen. 

Pats, p. 08 (1907). 
Pteropus (Spectrum) livingstouii, Matschie, Mcc/achir. p. 30(1899: 

Johanna) ; Trouessart, Cat. Mamm., Suppl. p. 54 (1904 : 

Pteropus edwardsi (pt., 7icc E. Geoff.), Peters, MP. Ahnd. Perlin, 

1867, p. 325. 

Diagnosis. — Allied to Ft. melanopogon, but cars semicircularly 
rounded off above, fur much longer, furred area of back of normal 
breadth, size smaller. Blackish above and beneath, with rump and 
sides of belly sprinkled with bright-coloured hairs, and with or 
without a tuft of tawny hairs on each shoulder. Forearm 162 - 
172 mm. Hab. Johanna I., Comoros. 

akuU and teeth.— Skull very similar iu general shape to that of 



Pi. melanoj^ior/on, but somewhat smaller, -with narrower preinaxillse 
and relatively smaller orbits. Upper and lower postorbital processes 
sometimes fused so as to completely encircle the orbits. — Structure 
of cheek-teeth as in the allied species, but m', m^, aud m^ relatively 
smaller. Vertical groove on front face of upper canines shallower, 
sometimes almost obsolete. 

Ears (dried skins). — Large, exposed ; differing in shape from 
those of any other species of this genus. Inner margin convex, 
outer margin sti'aight or flatly convex ; upper margin almost 
semicircularly rounded off. 

Wings. — Lateral membranes arising more closely together than 

Fur. — Much longer than in Ft. inelanopogon, rather harsh ; 
directed backward but not very closely adpressed on back, spreading 
on mantle. Approximate length on back 21, mantle 25-27, belly 
25 ram. Furred area of back not narrower than usual ; forearm 
and tibia nalced above. 

Colour. — Two adult skins, unsexed, teeth quite or almost unworr 

External measurements of Pteropus keyensis and livingstonei. 

Ft. keyensis. 
4 ad.* 

Ft. livingstonei. 

3 ad. 

(Incl. type.) 


Polles, total length, e. u 

,, metacarpal 

„ 1st jjhalanx 

2nd digit, metacarpal 

„ 1st phalanx 

,, 2nd -3rd jjbalanx, c. u. 
3rd digit, metacarpal 

„ 1st phalanx 

„ 2iid phalanx 

4th digit, metacarpal 

„ 1st phalanx 

,, 2nd phalanx 

6th digit, metacarpal 

„ 1st phalsEX ., 

,, 2nd phalanx 

Ears, length from orifice 

,, greatest width, flattened . 
Front of eye to tip of muzzle.... 

Lower leg 

Foot, c.u 











































18 t 












19-5 t 

* Type of species, and B.M., and Leyden Museum d and g. 

+ Estimate (skins). _ : , . 

rTERorrs ketensis and liyingstonei. 


Measurements of skulls and teeth of Pteropus keyensis and 

Skull, total length to gnathion 

,, palation to incisive foramina 

,, front of orbit to tip of nasals .... 
,, width of brain-case at zygomata. 

,, zygomatic width 

„ width across m', externally .... 

„ lachrymal width 

,, width across canines, externally . 

„ post orbital constriction 

„ iuterorbital constriction 

„ width of mesopterygoid fossa 

,, between p^-p^ internally 

„ between cingula of canines , 

,, orbital diameter 

Mandible, length 

, , coronoid height 

Upper teeth, c-m" 

Lower teeth, c-nij 

Upper incisors, combined width 

p', length 

,, width 

p-i, length 

„ width 

m' , length 

„ width 

m^, length 

„ width 

Pi, length 

„ width 

P3, length 

„ width 

P4, length 

,, width 

nj;, length 

,, width 

m^, length 

„ width 

m3, length 

,, width 

Pt. keyensis. 

Skulls': 6 ad. 

Teeth : 2 ad., 

2 imm.* 































Pt. livingstmiei. 
Skulls and 
teeth : 3 ad. 
(Incl. type.) 



72 1 


















75 1 




39 1 













* 6 ad., viz. type of species, and B.M., and Leyden Museum skulls 
d, g, h, i (Grand Key, Petit Key) 2 imm., tiz. Leydeu Museum, teeth of 
ekulls e,f. 

t Estimate. 


(type, and 88 5.9.1). Blackish or dark seal-brown above and 
beneath ; rump, sides of belly, and flanks more or less thickly 
sprinkled with golden ochraceoiis or tawny hairs, most of these 
hairs with concealed silvery greyish-white bases. "Tippet" (mantle) 
reduced to a tuft of rich tawny hairs on each shoulder (or one 
median tuft), the tawny colour confined to the tips of the hairs 
and shading through ochraceous-buff in subapical portion to buffy 
whitish in basal portion of hairs ; many of the blackish hairs in 
siioulder region tipped with golden tawny. 

A third skin (aged, unsexed, teeth well worn ; differs 
in having the mantle practically uniform blackish, with some of the 
hairs in the shoulder region tipped with chestnut. 

Measurements. On pp. 248, 249. 

Specimens examined. The British Museum material. 

Range. Only known from Johanna Island, Comoros. According 
to Humblot common in the large forests covering the summit of 
Johanna Island, but never seen in the low-lying portions of the 
island (Milne-Edwards & Oustalet, I. s. c). 

Type in collection. 

o. Ad. sk. ; skull. Johanna I., Comoros Earl Russell [P.]. 
{Dr. LiviiiffsUjiie). (Type of species.) 

i, c. 2 ad. sks. ; skulls. Johanna I. Purchased (Frank)., '2. 

G. The Ptbmopus raynehi group. 

Species. — Sis: Ft. cognutus, rayneri, ruhianus, Javellunus, yrandis, 
and chrysoprocius. 

Range. — Solomon Islands and Moluccas. 

General characters. — Rostrum shortened ; coronoid height of man- 
dible generally less than c-m.,. Posterior ledges of premolars short ; 
m^ reduced ; i^, i^, and Pj unmodified. Ears moderate, exposed ; fur 
of back short, adpressed; tibia hairy above (in Pt. chryso2}roctus very 
thinly so). Colour in one species nearly uniform brownish above 
and beneath, with paler mantle, in all the others the upperside is 
tricoloured (bright mantle, dark back, bright rump). Males of all 
species, except Pt. chrysoproctus, with well-developed neck-tufts. 
Size varying (forearm 120-177 mm.). 

Differentiation of species.— The extreme eastern species {Pt. 
cognatus : San Christoval) is externally (colour, size) rather similar 
to Pt. bru7ineus and Pt. lombocensis. In the other four Solomon 
Island species the rump is considerably brighter than the back, as 
bright as, or brighter than, the mantle ; this tricoloured style of the 
coloration of the upperside (bright mantle, dark back, bright rump) 
is found again in the Moluccan representative of this group, but in 
no other species of the genus. Pt. rayneri (Guadalcanar) is larger 
than Pt. cognatus; Pt. ruhianus (Ilubiana), Pt. lavelJanus (Vella 
Lavella), and Pt. grandis (Shortland and Bougainville) still larger. 
The fhiec latter species, which are closely interrelated, ditfer 


from each other in minor details of dentition, colour, and size. 
Ft. chryaojyr actus (Amboina group) is on the whole rather closely 
allied to tho largo western Solomon Island species. As yet no 
form of this group is known from New Guinea. 

Affinities of group.~Q\o&e\y allied to the Pt. hmhocensis group 
{infra), with which it accords in the shortening of the rostrum (a 
character still further developed in the hmhocensis group), reduced 
size of ffig, tendency in certain species to reduction also of m^, 
distribution of fur (tibia clothed above), and development of neck- 
tufts in males ; Pt. coynatus, in colour apparently the least 
specialized form of the group, shows, as mentioned above, 
externally no small resemblance to Pt. hmhocensis. The rayneri 
group differs from Pt. hmhocensis chiefly in the lesser shortening of 
the rostrum, the less constant and less conspicuous reduction of m^, 
and the perfectly normal i^. 

42. Pteropus cognatus, K. And. 
Ptcropus rayneri (pt.), Dobson, Cat. Chir. B. M. p. 33. 

Pteropus rayneri (pt.), Gray, Cat. Monk. SfC. p. 108 (1870 : 
San Christoviil ; nee Guadalcanar) ; Dohson, I. s. c. (1878 : 
San Ciiristoval) ; Trouessart, Rev. 4" Mag. Zool. (3) vi. p. 204 
(1879: San Christoval) ; id., Cat. Mamm. i. p. 78 (1897: San 

rtei-opus (Spectrum) rayneri (pt.), Trouessart, Cat. Mamm., SuppL 
p. 51 (1904: San Christoval). 

Pteropus cognatus, K. Andersen, Ann. <^- Mag. N. II. (8) ii. p. 365 
(1 Oct. 1908: San Christoval). 

Diagnosis. — Rostrum short and broad ; m^ conspicuously reduced 
in size. Ears moderate, exposed, narrowly rounded off at tip. 
Tibia clothed above. Dark brownish above and beneath (rump 
not differing from back) with cinnamon or russet mantle. Forearm 
at least \'2\ mm. (no jierfectly adult specimens seen). Hah. San 
Christoval, E. Solomon Is. 

/SYlm^Z.— Chief characters (compared with the typical Pteropine 
skull) : Rostrum short and broad, front of orbit vertically above 
posterior half of p' ; mandible heavier posteriorly : coronoid height 
(26 mm.) subequal to length of lower tooth-row, c-va^ (25-5 mm.). 

Teeth. — m^ considerably reduced, little more than half the size 
of Pj ; also m- slightly smaller than in Pt. liypomelanus. Cingulum 
of upper and lower canines stronger than in Pt. hyijomelanns, forming 
a well-defined rather broad ledge at inner base of teeth. Other 
teeth scarcely differing, neither in form nor in size, from those of 
Pt. Tiyj^omdanun. 

Fur. — Rather short, adpressed on back ; approximate length of 
hairs, back 11-12, mantle 14, belly 14 mm. Tibia thickly clothed 
above for proximal two-thirds or three-fourths, underside of tibia 
hairy for proximal half. 

Cohur. — (^ imm. skin, December, type of species: General 


aspect of back and rump Prout's brown ; individual hairs seal-brown 
at base, with short paler brownish or bufly brownish tips, in- 
completely concealing the darker base of the fur. Rump not 
differing in colour from back. — Entire underside approximately 
Prout's brown, of a rather paler tinge than back, palest (indistinctly 
washed with huffy russet) on foreneck, and darkening almost to 
seal-brown along centre of breast and belly. — Mantle and sides of 
neck cinnamon, slightly tinged with russet, becoming paler 
posteriorly at shoulders ; concealed base of hairs seal- brown. A 
tuft of buff-yellow glandular hairs on each side of neck, nearly 
concealed by surrounding darker fur (probably sexual character). — 
Crown and sides of head similar to back. 

A second specimen (male, a little younger than the above ; 
December ; skin) is slightly paler in colour, owing to the pale- 
coloured tips of the hairs being rather longer and more approaching 
huffy cinnamon. 

Measurements. On p. 2-57. 

Specimens ea-amined. The British Museum material. 

Mange. San Christoval, E. Solomon Islands. 

Ttipe in collection. 

History in literature. — Gray's Pt. rayneri (1870, I. s. c.) is a 
mixture of two species, the one from Guadalcanar (his " male " and 
" female "), the other from San Christoval (his " young ''). Since 
Pt. rayneri was based by Gray primarily on the adult " male " and 
"female" (which in fact are two adult females), both of which are 
from Guadalcanar, the name Pt. rayneri must be restricted to the 
Guadalcanar species. The characters of the San Christoval species 
were pointed out by me, and the species named Pt. cognatus, in 
1908 (L. s. c). The British Museum specimens of Pt. cognatus are 
the only ones on record (February, 1909). 

Pemarlcs. — Prom Pt. rayneri, with which this species accords in 
the characters of the skull and teeth (except perhaps in a still more 
conspicuous shortening of the rostrum), it is readily distinguished 
by having the rump similar in colour to the back, whereas in 
Pt. rayneri the rump is much brighter than and contrasting with the 
back. It is probably smaller than Pt. rayneri. Externally (dis- 
tribution of fur, colour, size) it bears much resemblance to the 
Australian species of the Pt. hypomelanus group, Pt. brunneits, 
which however is typical hypomelanine in skull and dentition. 

a. (S ad. skull. SanChristoval, Solomon Museum of Economic 

Is. ; Dec. 18.54 (J. Geology [P.]. 

b J' imm. sk. ; San Oliristoval ; Dec. Museum of Economic 

skull. 1845 {J. MacGilli- Geology [P.]. 

(Tyipe of species.) 

c. cJ juv. sk. ; San ChristOTal ; Dec. Lords of the Admiralty 

skull. 1854 {Br. F. M. [P.]. 


43. Fteropus rayneri, Gray. 
Pleropus 7-apieri (pt.), Dobson, Cat. Chir. B. M. p. 33. 

Pteropus rayneri (pt.), Grat/, Cat. Monk. ^c. p. 108 (1870: Guadal- 
canar; nee San Christoval) ; Dobson, I. s. c. (1878 : Guadalcanar) ; 
Trouessart, Eev. ^ Mag. Zool. (3) vi.p. 204 (1879: Guadalcanar) • 
id., Arm. Sci. Nat. (6) Zool. viii. Art. 12, p. 16 (1879 : remarks 
on distr.) ; Thomas, P. Z. S. 1887, p. 322 (Guadalcanar) ; id., 
P. Z. S. 1888, p. 472 (Aola, Guadalcanar) ; Trouessart, Cat. 
Matmii. i. p. 78 (1897: Guadalcanar). 

Pteropus (Spectrum) rayneri (pt.), Matschie, Mecjachir. p. 22 (1899 : 
Aola, Guadalcanar) ; Trouessart, Cat. Mamm., Supnl p 51 
(1904: Guadalcanar). ^' 

Diagnosia. — Similar to Pt. cognaius, but rump much brigliter than 
and contrasting with back. Eorearm 137-5-140-5 mm. ITab, 
Guadalcanar, E. Solomon Is. 

Skitll and teeth.- — Skull similar to that of Ft. cognatus, though 
perhaps slightly larger, and with the rostrum not quite so much 
shortened (compare measurements, p. 257). Teeth scarcely 
differing from those of the allied species. 

Palate-ridges (two specimens; S8.1.5.2, 3).— N'o special modi- 
fications. Formula 5 + 5 4- 3. First ridge terminating laterally at 
front of canine ; second at back of canine ; third at front of p' • fourth 
at back of p^ ; fifth at middle of p^; sixth at front of m^ ; seventh 
at hack of m' ; eighth at back of m^ ; ninth and tenth behind m'- ; 
eleventh to thirteenth situated at palation border. 

Colour. — 5 ^^- skin, teeth slightly worn, December; cotype 
of the species ( : — Back uniform seal-brown with a slight 
purplish gloss, rather more inclining to vandyck-brown laterally 
along membranes, and posteriorly near rump. Kump and upperside 
of femur much brighter than and contrasting with back, buffy clay 
washed with ochraceous-buff anteriorly near back, and shadino- to 
mars-brown on outer side of femur along membrane and upperside of 
tibia. — Breast uniform seal-brown. Flanks vandyck-brown, shading 
to mars-brown on belly, anal region, and underside of femur and 
proximal part of tibia. — Mantle and occiput deep cinnamon-rufous 
approaching rich russet, contrasting with seal-brown back, passing 
through a darker shade on sides of neck, to vandyck-brown on .fore- 
neck, the colour of this latter being scarcely different iu tinge from 

that of the flanks. Concealed base of fur of mantle seal-brown. 

Crown between Prout's brown and mars-brown, passing throuo-h a 
darker shade on sides of head to almost seal-brown on chin and 
throat, the colour of this latter very similar to that of the breast ■ 
crown, sides of head, cliin, and throat sparsely sprinkled with silvery 
white hairs. 

In the second cotype (same locality, sex, age, and month ; skin ; the contrast between mantle and back, and back and 
rump is less striking, owing to the darker tinge of the mantle and 
rump : mantle approaching burnt-umber, rump pale russet with a 
slight wash of clay. Underside as in the foregoing, but the seal- 
brown colour restricted to the centre of the breast ; sides of breast 
fianks, belly, and anal region uniform p.Tlc vandyck-brown. 


The amount of palo (silvery white, cream-buff}', or pale buff}) 
sprinkling of the fur of the head is individually variable ; in one 
specimen ( $ irani. al. ; the pale hairs are so much in 
excess of the dark ones as to make the whole of the face, including 
the chin, cream-buffy, strongly contrasting with the mantle, back, 
and underparts. 

ISe.vwil differentiation. — Males differ from females in having 
a tuft of rich orange-buff glandular hairs on each side of the neck, 
nearly concealed by the surrounding darker fur. Tlie tuft ia 
present also in immature males, but absent in three females 

Measurements. On pp. 256, 257. 

Specimens examined. The British Museum material. 

Range. Guadalcanar, E. Solomon Islands. 

Coti/pes in collection. 

History in lilerature. — Gray's Pt. rai/neri (1870, I. s. c.) was based 
primarily on two adult skins collected by J. MacGillivray and F. M. 
Rayner in Guadalcanal*. These two specimens (described by Gray 
as " male '' and " female," but in fact both of tliem females) must 
therefore be taken as cotypes of Pt. rai/neri, and Guadalcanar as the 
type locality of the species. Two immature specimens, obtained by 
MacGillivray and Rayner in San Christoval, and described by Gray 
(l. s. c.) as the "young" oi Pt. rai/neri, are a distinct species, 
Pt. cognaius (antea, p. 251). — The cotypes of Pt. rciyneri (and an 
odd skull in the British Museum, probably also dating from 
MacGillivray or Rayner) remained for many years the only speci- 
mens known, until in 1887 C. M. Woodford collected a few 
examples in Aola, Guadalcanar, three of which are now in the 
British Museum, one in the Dresden Museum. So far, these are the 
only specimens on record (February, 1909). 

Remarlcs. — Externally this species is readily distinguished from 
Pt. cognatus by its bright-coloured rump. In specimens which 
have this character strongly pronounced, the upperside is strikingly 
tricoloured: mantle some shade of russet, back seal-brown, rump 
buffy clay, the rump being decidedly brighter than the mantle, and 
the contrast between rump and back therefore greater than between 
mantle and back ; this tricoloured pattern of the upperside is 
found again in Pt. rubianns (New Georgia group), Pt. lavcllanus 
(Vella Lavella), Pt. grandis (Bougainville group), and Pt. chrgso- 
proctus (Moluccas), but in no other known species of the genus. 

a. Imm. skull. Not reg. 

b. 5 ad. sk. ; Guadalcanar, Solomon Museum of Economic 

skull. Is.; Dec. 1845 (./. Geology [P.]. 


c. $ad. sk. ; Guadalcanar; Dec. Lords of the Acliuiralty 

skull. 1854 {Di: F. M. [P.]. 

Rayner). (b, o: coft/pes of speaies.) 

il-f. S iram., S Aola.'Guadalcariar. C. M. Woodford, Esq. 

ad., 2 [C.]. 
imm., al. ; 

skull oT C. ;. ■ . ,. , 1 > . .1 •.. r : . i 

prnropus r.LiUAXtJs. 2.35 

44. Pteropus rubianus, A". And. 

rter.jpus grandis (pt.), Thomas, P. Z. S. 18S8, p. 470 (1889: 
Kubiana, Solomons); Trouessart, Cat. Mamm. i. p. 80 (1897: 
Solomons) ; MaUchie, Megachir. p. lo (1899 : Rubiana) ; 
Trouessart, Cat. Mamm., Suppl. p. 49 (1904: l\ubiana). 

Pteropus rubianus, K. Andersen, Ann. &■ Mag. N. H. (8) ii. p. 366 
(1 Oct. 1908: Rubiana). 

Diagnosis.- — Allied to Ft. rayneri, but much larger. Back 
vandj-ck -brown, rump strongly contrasting j-ellowish buff, mantle 
and foreiieck dark russet, sliading to deep tawny on sides of breast 
and belly, and to yellowish buff on crissum ; centre of breast seal- 
brown ; forehead and sides of face mottled yellowish buff and 
chestnut. Forearm 1G3 mm. Hah. New Georgia group, Central 
Solomon Is. 

ShulJ. — Scarcely differing in general shape from that of Pt. 
rayneri, but much larger and more heavily built, -with strong 
sagittal crest, and unusually deep zygomatic arches. Rostrum 
rather short, deep, distiucfly compressed laterally ; front of orbit 
vertically above front of m\ 

Teeth. — Structure of teeth essentially as in Pt. raipieri. Upper 
canines heav}' at base, antero-posterior basal diameter 5 mm. ; 
ciugulum strong, forming a well defined, rather broad ledge ; crown 
in profile faintly recurved, vertical groove on front face deep and 
broad ; vertical Iceel on lingual face sharply projecting, p' a minute 
spicule. Posterior ledge of p^ and pg short, but distinctly marked 
off from tooth postcro-externally. Ledge of p'' and p^ very short, 
postero-external notch distinct though very small in p\ obsolescent 
in p^. m^ much smaller than p^, slightly larger than m^. — i, twice 
and a half the bulk of ij. Lower canines heavy at base : cingulum 
forming a broad ledge ; crown recurved. Pj large, twice the size 
of ij. m^ very small, subequal to i^. 

Palate-ridges (one, the type). — 5 + 5 + 3. Arrangement as in 
Pt. rayneri. 

Ears. — Short, covering about half the distance from base of ear 
to back of eye, when laid forward. Outer margin flatly concave in 
upper third, tip of ear ralher narrowly rounded off. Basal half of 
hinder face of conch hairy. 

Wings. — Membranes about 20 mm. apart at origin from back. 
Interfemoral. — Short in centre ; depth about 7 mm. 
Fur. — Hair of back rather short, adprcsscd. Approximate 
length, back 13, mantle 19, belly 15 mm. Least width of furred 
area of back about 47 mm. 

Above, fur extending in a narrow line of short closely adpressed 
hairs along humerus ; region round elbow naked ; forearm naked, 
except for some short thinly set hairs on proximal third. Femur, 
knee, proximal three-fourths of tibia, membrane along outer side 
of femur and proximal half of tibia, and interfemoral along 
inner side of femur and proximal three-fourths of libia, thickly 
furred. — Below, forearm and tibia pruclicalh- naked. Antebrachial 



membrane, lateral membrane along outer side of forearm aud be- 
tween humerus and femur covered with rather long wooUj- hairs. 

Colour (type, (J ad. al.).— Back glossy brownish, between 
vandyck-brown and burut-umber, with a tinge of goldeu butfy, this 
colour extending backward in a narrow line along sides of rump 
next to membranes. Rump, upperside of interfemoral, and thighs 
yellowish huffy, much lighter than mantle and strongly contrasting 
with back ; upperside of tibia3 similar, but washed with ochraceous ; 
base of fur of rump seal-brown. — Centre of breast dark seal-brown, 
forming a large well-defined patch. Sides of breast, flanks, and 
belly rich tawny, this colour conHned to tips of hairs; sub- 
apical portion of hairs paler, ochraceous or ochraceous-buff, extreme 
base seal-brown. Anal region and thighs yellowish buff, nearly 
similar in tinge to rump. — Occiput, mantle, sides of neck, foreueck, 
and throat rich glossy russet, approaching pale chestnut, darker 
than sides of breast, and lightening to yellowish ochraceous-buff on 
shoulders ; extreme base of hairs of mantle dark brown. A tuft of 
rigid, orange-ochraceous glandular hairs on each side of neck, 
contrasting in colour with surrounding darker fur (sexual character). 
Forehead and sides of face mixed yellowish buff and chestnut, 
shading on crown gradually into bright chestnut of occiput. 

External measurements o/Pteropus rayneri and rubianus. 

Pf. rayneri. 
3 ad. 

(Incl. cotypes.) 

Pi. ruhianus. 
S ad. 

MiN. Max. 

mm. mm. 
137-5 140-5 
57 63 
11-8 128 
29 34-2 
07 70 
16-7 18-7 
132 15-2 
89 92 
68 69 
96-5 101 
85-2 90-5 
54-2 58-5 
55 65-5 
92-5 96-8 
40-2 43-7 
41 41-5 

59-2 63-5 





















PoUex, total leiio-th, c. u 

„ 2nd-3rd phalanx, c. u. ... 
3rd digit, metacarpal 

5th digit, metacarpal 

,, 1st phalanx 

I'lKUOl'lS (;'OGXATU>< ETC. 


Meas'd'Oiients of sJadls and teeth 0/ Pteropus cognatus, raj'iicri, 
lotd nibiaims. 

Pt. cognatus. 

I Skull: 1 ad. 

Teeth : 1 ad.. 

Vf. rayneri. 

Skulls: 2 ad. 

Teeth : 2 ad., 

1 iinin. 

(Incl. type.) -(Incl. cotypes) 

MiN. Wax. 

mm. mm. 

Skull, total length to gnatliion 

„ palation to incisive foramina 

„ front of orbit to lip of nasals 

,, width of brain-case at zygomata... 

,, zygomatic widtli 

,, width across m', e.\lemally 

„ lachrymal width ... 

,, width across canines, externally... 

,, postorbital constriction 

,, interorbital constriction 

,, width of mesopterygoid fossa 

„ lietween p^-p'', internally 

„ between cingula of canines i 

,, orbital diameter I 

Mandible, length ! 

„ coronoid height 

Upper teeth, c-m^ 

Lower teeth, c-m, 

Upper incisors, combined width , 

Upper canines, ant.-post. basal diameter ., 
Lower canines, ant.-post. basal diameter .! 

p', length ] 

„ width 

p*, length I 

,. widtli i 

m', length | 

width 1 

, length I 

width I 

Pj, lengtii 

„ width 

Pj, length 

,, width 

p^, length 


in ,, length 


iBj, lengtii 


,. length 






























Mm. Ma.\. 

mm. nun. 



























Pt. ruhianus. 




































Measurements. On pp. 256, 257. 
Specimen examined. One, the type. 

llaiu/e. As yet kuowii only trom Eubiana, off Xow Georgia, 
Central Solomon Islands. 
Tiipe in collection. 

a. rf ad. fil. ; iikull. Eubiana, Centi-al C. 11. Woodford, Esq. 
Solouion Is. [C.J. 

( Ti/'pe of sperips.) 

45. Ptei'opTis lavellanus, A'. And. 

Pteropus lavellaniis, K. A7idersen, Aim. ii- Mikj. N. II. (8) ii. p. 3(50 
(1 Oct. 1908 : Vella J.avella, Solomon Is.). 

Diagnosis. — Allied to Ft. rvbianus, but smaller, with much 
shorter tibia, and much darker colour of the fur. Crown and face 
griz/led buffy, greyish, and dark brown. Forearm 151-155-5 mm. 
Hah. Vella Lavella, Solomon Is. 

Skull and teeth. — Skull distinctly smaller than in Pt. mhianu?, 
hut scarcely differing in shape. Dentition as in Pt. rnbianus, but 
ciiigulum of upper and lower canines broader than in any other 
form of the Pt. rayneri group. 

Pi,r. — Length "and distribution of fur as in Pt. ruhianw^. 
Apiiroximate length, liack 13-15, mantle 18-21, belly 17-18 mm. 
Least width of furred area of back about 45 mm. Tibia thickly 
haired above for proximal three-fourths. 

Colour. — Type, S ad. skin, teeth well worn : — Back glossy seal- 
brown, this colour continued backward in a narrow line along sides 
of rump next to membranes and on upperside of tibiae. Enmp and 
inner side of thighs and tibias pale cinnamon-rufous, paler than 
mantle and strongly contrasting with dark back ; base of fur of 
rump seal-brown. — Centre of breast and upper belly glossy 
blackish, forming a large oval patch ; sides of breast and belly, 
including flanks, dark mars-brown, contrasting with blackish 
pectoral patch, and shading gradually into dark cinnamon-rufous on 
crissum. — Occiput, mantle, sides of neck, and foreneck chestnut- 
chocolate ; extreme base of fur blackish. A tuft of rigid, orange- 
tawny glandular hairs on each side of neck, on spreading con- 
trasting in colour with surrounding darker fur (sexual character). 
— Crown and intcrocular space similar to occiput, but conspicuously 
spriukled with tawny and buffy hairs. Circumocular space and 
sides of face mixed blackish, buffy, and pale greyish. Chin and 
throat chestnut seal-brown. " Iris brown " (A. S. llcek). 

In three sjiecimens examined the individual variation is chiefly 
confined to a brighter or darker tinge of the rump and a greater 
or smaller admixture of pale hairs in the colour of the crown 
and face. Rump varying from cinnamon-rufous, through cinnamon, 
to nearly orange ocliraceous-buffy. In the palest-headed specimen 
( imm.) the crown and forehead are golden buffy sprinkled with 
prevish-white and blackish hairs. 

PTF.llOHL'^; GliAXUIS. 2o'.J 

St'.vual dlffereiitiiitioii. — A tuft of rigiJ, biighler-colouied 
gliiudular haiis on each side of neck in males, wautiug in females. 
Canines heavier in males than in females. 

Measurements. On pp. 2(33, 264, 

iSpecimens examined. Three, as registered below, 

liange. So far only known from Vella Lavella, Xew Georgia 
group, Central Solomon Islands. 

I'^ipe in colled ion. 

liehiarln. — This species differs from Pt. ruhiwnis chiefly in (he 
characters given in the diagnosis above. From Pt. r/randis, which 
it closely resembles in the colour of the neck, back, rump, and 
Tiiiderparts, it is readily distinguished by its smaller size, relatively 
shorter tibia and smaller ears, and by having the crown and face 
grizzled buffy, greyish, and dark brown, not uniform blackish or 
seal-brown as in Pt. grandis. 

a. J u(l. sk. ; skull. Vella Lavella, SolomonB ; A. S. Meek 8.11. 1«.2. 
12 March, 1908. [C.]. 

{Type of species.) 
A.r. 9 iiiim., 2acl.sks.; Vella Lavella ; 17 Feb., A. S. Meek 
skulls. 2 March, 1908. [0.]. 

4ti. Pteropus grandis, Thos. 

I'teropus grandis, nomas, Ann. iff Mni/. i\. II. (5) xix. p. 147 (1 Feb. 
1887 : Shortland I.) ; id., P. Z. S. 1887, p. 320, pi. xxv. (animal), 
text-tig. 1 (teeth) (Ala, Shortland 1.) : Woodford, Not. amom/ 
the Head-liunter.<, p. 197 (1890: Shortland grou])) : Troiiessart, 
Cat. Mamm. i. p. 80(pt.) (1897 : Solomons) : Matachie, Megachir. 
p. 15 {pt.)(1899 : Alu) ; Trouessart, Cat. Mamm., Stippl. p. 49 (pt.) 
(1904 : Shortland) ; Mdkr, Fam. S,- Gen. Bats, p. 58 (1907). 

Diagnosis. — Similar to Pt. Javellanus, but larger with relativelv 
longer tibia and larger eary. Crown and face uniform blackish 
seal-brown. Forearm 167-1T2 mm. JIuh. Bougainville grouj), 
Solomon Is. 

Colour. — Type, e? ad, skin, Alu : — Back glossy blackish seal-brown, 
this colour continued backward in a narrow line along sides of 
rump next to membranes and on upperside of tibia?. Paimp pale 
ochraccous-buff, much paler than mantle and strongly contrasting 
with back, shading into cinnamon-rufous along inner sides of 
thighs, and this in turn into chestnut along inner sides of tibia>. 
Base of fur of rum]) seal-brown. — Centre of breast and u])per belly 
glossy Vjlackish, forming a large patch or broad stripe ; sides ot 
breast and belly, including Hanks, dark chocolate, shading gradually 
into dark cinnamon-rufons on crissum. — Occiput, mantle, sides ot 
neck, and foreneck dark chocolate, similar to sides of breast and 
not very strongly contrasting with back. .V tuft of rigid, mars- 
brown glandular hairs on eacli side of neck, on spreading somewhat 
contrasting in colour with surrounding darker fur (sexual character). 
Crown, forehead, sides of face, chin, and throat blackish. "' Iris 
yellow-brown'' (V. M. \\'oodford). 

An adult male from Bougainville (skin, 8.] 1.16.4) is similar to 


the Jbregoiug, l)ut nini]) still brighter, nearly buft-ycllovv, with 
extreme base of hairs seal-brown. " Iris browu '" (A. S. Meek). 

Se.rrud diff.retUintwn. — Probably as in Pt. lavcUamw (p. 259), 
but DO females seen. 

Measurements. On pp. 203, 264. 

Specimens e.ramined. Three, as registered below. 

Itawje. Shortland Island (Ahi) and Bougainville, W. Solomon 

Type in collection. 

llcniarlcs. — The differential characters given in the diagnosis of 
Ihis species rest on a cosnjtari.son with J^t. hivdlinms. From 
Pt. riibianus it is easily discriminated by its larger ears and much 
darker colour. Tl;e vandyek-brown tinge of the back in Pt. ruhianua 
is in Pi. grandis rei)laced by blackish or dark seal-browu ; tbe dark 
russet and deep tawny of tlie mantle, foreneck, and sides of breast 
and belly, by dark chocolate ; the mixed huffy and chestnut of the 
face, by blackish. The colonr of tbe fur of Pt. rp-andis is, on the 
whole, mnch like that of Pt. lai'cUanus. except for the complete 
suppression of tbe light sprinkling of the colour of the face, so 
conspicuous in Pt. JareUanvs. In the characters of the skull and 
teeth Pt. ip-andis is similar to Pt. rnhiamis ; cingulnm of canines 
as in that species, narrower than in JH. lavellannx. The lengtli and 
distribution of the fur is practically tlie same in all three species. 

ff. Ad. skull. Aln, ^;llortlam^ I., C. M. Woodford. Esq. 

Hohiiiions ; April, [C.J. 


i'. c? ad. &!;.: skull. Alu ; April, 1S86. C. M. AVoodford, Esq. 

[C ]. ( Type of species.") 

c. ci ftd. fk.; skull. Ciiin, BougainvilU>, A. 8. Meek |C!]. 8.11.1fi.4. 

SoUiiiioiis ; 11 •T:iu. 


-17. Pteropus chrysoproctus, Tuiuu. 

PUroims elii->i!>opi-octiis (pt.), Dobson. Cat. Chir. ]!. il. p. 47. 

I'tcropu.i chrysoproctus, TemviiinI;, Moii. Mtnnm. ii. p. fi", pi. .\.\xv. 
ii^:. 2 (head), pi. xxxvi. tigs. 7, s (i-kull) (1K57: Amboina) ; 
IVafjncr, Sc/nrher'K S(iu(/., ib'ujjjL i. p- •■i4S (1839: Amboina); 
<S'. AliU/er, in 'J'nnminrk'.f Nat. (jesch. Arderl. Ocer~. L'cz.. Zont/d. 

PJ\'..J(l./ , ._.....y, t........ .'v.-.-^v. ^.-.^ , . .... ^.. .> 

yxurtc/. ,^...«w...... , Wdi/nfr, Svltrfhcr !< SiiiKj., Snppl. v. p. o07 

(IHoii-oi) : Amboina^; (Icrvnu, Ilist Nut. Mfniim. i. )>. 187 
(18-54: Anib.jr.a) : Giehel, >>aw/. p. i'!'4, footnutc (ISo.J; .Am- 
boina); I'inseh, Neu-(ruinea, p. 15(l(18Ho: .Ainboina); Peterii, 
MB. Aki'.il llrr/in, ]8r>7, p. ."Wl (Aiiiboira; Ocvnni; I'ulo IVi- 
jang; Ciovaiii); Fita'n(/('i; Sli. Ahail. IVifi, l.\. Abtli. i. ]). 425 
(1870: Aiiiboina) ; Mur<-hi, Atti S,><: Itat. Sri. .\,it. xv. p. 515, 
')1. viii. fiir. 2 (1872-73: structure of bail's): Dobgon, Cat. Chir. 
'U. M. p. 47 (pt.) (b^7S : Waluljelbi I.) ; Po^iuh'nj, .Muhiij. Arch. 

PTDKoPl'S (■IIUVm)IM:(J( TtlS. 261 

p. .S22 (1878: Ceram) ; Truucssart, Rev. ^ Ma//. Zool. (.">) vi. 
p. 205 (1879: Amboina ; Ceiaiu ; T. Panjan^ ; Goram ; Watu- 
bella); Jentinlc, Cat. Ost. 3Ia»im. p. 2.57 (1887: Amboira ; 
Ceram ; Keffing' I.; P. Panjang; Goram; Saughir) ; ?a., Cat. 
Sijst. Mamm. p. 144 (1888 : Amboiua ; Ceram ; Arsilulu ; Keffing ; 
P. Panjnng; (-ioram ; Sangbir) ; Trouessart. Cat. Mamm. i. p. 80 
(pt.) (1S97 : Amboina; Ceram ; P. Panjaiig; Goram ; Watiibella); 
Matxchie, Mi'yachir. p. 14 (pt.), pi. ix. figs, o, 5 r?, 5t (skull) 
(1899: Amboina; Ceram; Goram); Trouessart, Cat. Mannn., 
6'it»;>/. p. 49 ( 1 904 : Sangbir; Amboina; Ceram; Goram ; Watu- 
bella) ; Willink, Nat. TIjd. Nerlerl. Iiid. Ixv. p. 272 (pt.) (1905 : 
Amboina; Ceram; Goram); Miller, Fain. &■ Gini. Bats, p. 58 
Kunvcteris pbaiops (pt., nee Ptero/>u,s phaiops, Temni.), (ji-ai/, Cat. 
Monk. S,-c. p. 113 (1870 : Watubella 1.). 

Diagnosis. — Allied to Pt. gramfis, but canines slenderer nt 
base, i., and p, smaller, cbeek-teeth narrower, np])erside of tibia 
more sj'ur.'sely haired, colour of fur paler. Back dark brownisb, 
rump similar or much paler ; breast and belly some shade ol 
cinuamon-rufous or chestnut or seal-brown, rarely uniform, 
generally distinctly darker in centre than on sides of breast and 
belly ; head and neck all round tawny or tawny ochraceous, con- 
trasting Avith darker back, ilalcs without bright-coloured neck- 
tults. Forearm l(i3-17'j"r) mm. I/ab. Amboina grouj). 

Skull. — Size and general characters as in Pt. rubiniius and 
(frandis, but orbits slightly larger, rostrum longer ; orbital diameter 
lo-lo'o ram. (l;5 8-l4'S in Pt. ruhi.arnis aiul qivdidis), rostruin 
from front of orbit to tip of nasals 20-29 (22'6-24), palate from 
palation to incisive foramina 87-39 (34'."i-3(3"2). 

Ti'tth. — Upper incisors smaller than in Pt. rubiauns and (/randltt. 
Canines above and belovr practically of the same height as in the 
allied species, but considerably slenderer at base and with much 
narrower cingula ; antero-posterior basal diameter of upj)er canines 
4-4*8 mm., against 5 5'0 in Pt. ruhianns and grandis; of lower 
canines 3-3-8, against 4-4"5. i., and p^ conspicuously smaller ; 
i., once and a half or nearly twice (in l^t. rnlnanas and graiiditi 
twice and a half or nearly three times) the bnlk of ij ; crown of ]\ 
2'o-28 X 2-2-2"4 mm. (in Pt. ruhianun and grandis 2-8-3-3 x 
2'7 ;'))• i)'-m' and p^-na., scarcelj- shorter than in the allied 
species, but considerably narrower; see detailed measurements 
p. 2(54. 

Fur. — Length and distribution of fur essentially as in Pt. ruhianus 
and grandis, but upperside of tibia thinly bailed or naked. 

Colour. — cT ad. skin, Burn, teeth somewhat worn, 97. 12. H.I : — - 
General colour of hack glossy dark burnt-nmbor. Individual hairs 
tawny or tawny cinnamon-rufous at base, seal-brown at ti]). 
Owing to a gradual increase, in direction from front of back to 
lump, in the length of the pale-coloured bases of the hairs and 
corresponding decrease in the length of the dark tips, the colour 
becomes more and more distinctly tinged with golden tawny on 
po-;ieiior portion of back near ibis latli^' tinge again shading 

2G2 riEUorrs ciiuYsorRncTL's, 

into tawny ocliraceous on rump and into nearly ochraccous-ljuff on 
thighs. The colour of the rump and thighs thus becomes similar to 
or rather lighter than that of the mantle, and strongly contrasting 
with dark front of hack. — Breast, bellj', and flanks dark russet, 
darkest (washed with chocolate) on centre of breast, palest (tinged 
with huffy) in anal region ; base of fur everywhere paler, ochra- 
ceons, ochraceous-bufF, or buffy. — Head, mantle, sides of neck, and 
foreneck nearly uniform tawny ochracenus. 

d yg. ad. skin, Ceram, Differs from foregoing chiefly 
in the following particulars : — Back darker, glossy seal-brown ; 
rump and thighs similar, but slightly washed with chocolate. A 
narrow longitudinal stripe of deep maroon seal-brown along middle 
of breast and belly, extending orer anal region and underside of 
thighs ; sides of breast and belly, and flanks huffy cinnamon-rufous. 
Circumocular space, sides of muzzle, chin, and throat seal-brown. 

5 ad. skin, Ceram. teeth slightly worn, : Differs from 
the Burn specimen only in tlie colour of the breast, belly, and 
flanks. Centre of breast glossy blackish tinged with maroon, 
gradually shading into chestnut on sides of breast and flanks, and 
into chestnut cinnamon-rufous on belly and anal region. 

c^ ad. skin, Watubella I., teeth slightly worn, 
Similar to the young adult male from Ceram (7.1.1 .244) in the colour 
of the head, back, rump, mantle, sides of neck, and foreneck, but the 
whole of the breast, belly, flanks, and anal region glossy seal-brown, 
tinged with chocolate on sides of breast and belly ; base of fur of 
sides of breast and belly ochraceous. 

The differences in colour described above appear to be independent 
of locality, sex, and age. The four cotypes of the species (Amboina) 
in the Leyden Musetim and a topotype in the Berlin Museum show 
somewhat similar individual variations, though none has the whole 
of the underside of the body so uniformly dark-coloured as the 
Watubella specimen. The variations, in the whole series examined, 
may be briefly epitomized as follows : — Back glossy seal-brown or 
washed with a j)aler shade of brown approaching dark burnt- 
umber; rump and thighs sometimes similar to back, but inmost 
specimens much paler (similar to mantle) and contrasting with back. 
Breast, belly, and flanks sometimes nearly uniform dark russet with 
paler bases to the hairs (palest-bellied extreme), but generally with 
some seal-brown colour on centre of underparts, this dark tinge 
sometimes confined to centre of breast, sometimes extending also 
to centre of belly and anal region, rarely uniformly covering the 
whole of the breast, belly, anal region, and flanks (darkest-bellied 
extreme). Head, mantle, sides of neck, and foreneck some shade 
of oohraceous-huff or tawny ochraceous ; circumocular region, sides 
of muzzle, chin, and throat similar to crown or darkened with 

Sexual differentiation. — Canines conspicuously longer and slightly 
heavier in males than in females. Measurements taken on three 
males and one female : upper canines, length from alveolar border 
10-2-10*8 (males) and (female), antero-posterior basal diameter 

PTKUOPus ciiKYsoriiocrus. 


4'3-4*8 (males) and 4 (female) ; lower canines, length 8-9 (males) 
and 7"5 (female), anteio-posterior basal diameter 3'2-3-S (males) 
and 3 mm. (female). (J^o distinct rigid and brightttr-coloured 
glandular neck-tufts detectable in males, nor in females.) 

Measurements. Below and on p. 264. 

Specimens e.vainiiied. Ten, in the collections of the Leydcn 
(specimens /, r/, h, l ; Amboina), Berlin (nos. 2626 and 3318 : 
Amboina and Ceram), and British Museums, including the cotypes 
of the species. 

Ran(je. Amboina group : Buru, Amboina, Ceram, Arsilulu, 
Keffing, Pulo Panjang, Goram, Watubella. The species has been 
recorded in literature from the Saiighir Islauds (Siao) ; see 
synonyms, pp. 260, 261. 

Cotypes in the Leyden Museum. 

Pteropus chrysoproctus, Temrainck ; 1837. — Type localitj', Am- 
boina ; cotypes, four mounted specimens (originally in alcohol), 
collected by Miiller and Macklot, specimens /, g, h, and i of 
Jeiitink's ' Catalogue systeroatique.' — The skull figured in ' Mega- 
chiroptera des Berliner Museums ' (l. s. c.) is that of an adult male, 
skin mounted, collected in Ceram by Wallace, Beg. no. 3318. 

liemarks. — This species is readily distinguished from the closely 
related Ft, ruhianus and grandls by the characters of the skull, 

External measurements o/ Pteropus lavellanus, grandis, and chrysoproctus. 

Pi". lfivM(inui<. 

2 ad. 

(Incl. type.) 

MiN. Max. 

! mrn. mm. 

Forearm ! l.')l 155'5 

Pullex, total length, c. u ! 66 69 

,, metacarpal 14 \b 

„ 1st phalanx 3.S-.5 34 

2nd digit, metncarpjil 745 78 

I ., Ibt phalanx 2n-.'> 21 

2nd 3rd phalanx, c. . I. . 16-5 19 

3rd digit, metacarpal 100.^ 107 

1 ,, 1st phalanx 77'5 78-5 

• ,, 2nd phalanx , 110 115 

I 4th digit, metacarpal ' 97 102 

Istuhalanx 6.S 63 

2ndpiialnnx \ 63 67 

5th digit, metacarpal 102-5 108 

1st phahinx 44 46 

,, 2nd phalanx 45 485 

Fars, length from orifice ' ?26» ?27 * 

Lower leg ' 65"5 67'5 

Koot.c. u 48 48-5 

Pf. c/randis. 

2 ad. 
(Incl. type.) 

Pt. chri/soproc/us. 
4 ad. 








15 5 
115 5 



















Mix. Max. 




















91 . 


1 24 



* Kstimate (skins). 



JHeasurc)}iC)i(s of slid Is and teeth 0/ Pt. lavellauus, graiidis, and chrysoproctus. 

Pf. lavellanii». 
Skulls: 2 ad. 
Teeth: 2 ad., 

1 ill! in. 
(Incl. tjpe.) 

Vt. graiidis. 

3 ad. 
(Incl. type.) 

P/. chryno- 


4 ad. 



Skull, total length to gnatliion fjX.Q 

., palation to incisive foramina '>n-"S 

,, front oC orbit to tip of nasals ^>.- 

„ ■width of bi-ain-fase at zygomata ... ~ 
zygomatic width 






,, width across ni', externally I, 

„ lachrymal width 

„ width across canines, externally 

,, postorbital constriction 

., interorbital constriction 

„ width of mesoplerygoid fossa ... 

,, between p'-p*, internally 

,, between cingnla of canines 

„ orbital diameter .„.^ 

Mandible, length ' ^^'r 

coronoid height 

Upper teeth, c-m''' 

Lower teeth, c~m^ 

Upper incisors, combined width 

Upper canines, ant. -post. b;isal dianieter 
Lower canines, ant.-post. basal diameter 

p', length 

„ width 

p*, lejigl h 

.. width 

m', length 

,, width 

ni'^, length ■■■■■ 

„ width 

pi, length 

,, width 


width ' 




















: li'i. length [ •^_^ 

i ., ^vidth I "g 

m^, length * - 

„ width Yc 

mj, length |'6 

„ width 1'^ 





















Mis. M.^x. Min. 








i 2-2 

I 2-8 

I 2-7 

i 5-2 

' 33 


! 3-7 

I 6-7 






36-2 ' 

24 1 
27-5 I 
42 5 

21-8 ! 

16 : 


8 I 

9 ! 

8 8 ' 

14-8 I